GLOBAL
BIODIVERSITY
STATUS OF THE EARTH'S LIVING RESOURCES
COMPILED BY
WORLD CONSERVATION MONITORING CENTRE
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Global Biodiversity
Status of the Earth's Living Resources
A Report compiled by
the
World Conservation Monitoring Centre
Editor: Brian Groombridge
WORLD CONSERVATION
MONITORING CENTRE
in collaboration with
The Natural History Museum, London
and in association with
IUCN - The World Conservation Union
UNEP - United Nations Environment Programme
WWE - World Wide Fund for Nature
and the
World Resources Institute
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NATURAL exh ©
MUSEUM UNEP wwe
With project sponsorship from
Overseas Development Administration, UK
and additional support from
The Ministry of Foreign Affairs, The Netherlands
The Ministry of the Environment, Denmark
and
The World Bank
iH
CHAPMAN & HALL
London e Glasgow e New York e Tokyo e Melbourne e Madras
1992
Published by Chapman & Hall, 2-6 Boundary Row, London SE1 8HN
Chapman & Hall, 2-6 Boundary Row, London SE1 8HN, UK
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This report is a contribution to GEMS - The Global Environment Monitoring System
First edition 1992
© 1992 World Conservation Monitoring Centre
Reproduced from camera-ready copy prepared by WCMC.
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the Martins Printing Group.
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The publisher makes no representation, express or implied with regard to the accuracy of the information
contained in this book and cannot accept any legal responsibility or liability for any errors or omissions that
may be made.
Citation: World Conservation Monitoring Centre (1992) Global Biodiversity: Status of the Earth's living
resources. Chapman & Hall, London. xx + 594pp.
Also available from IUCN Publications Services Unit, 181a Huntingdon Road, Cambridge,CB3 ODJ
Cover Photos Mugger, Crocodylus palustris: Brian Groombridge
Guzmania lingulata: D. Muleax
Fish market, Indonesia: Tom Moss/WWFEF Photo library
Henri Pittier National Park, Venezuela: Paul Goriup
The designations of geographical entities in this book, and the presentation of the material, do not imply the
expression of any opinion whatsoever on the part of WCMC or its sponsoring organisations concerning the
legal status of any country, territory, or area, or of its authorities, or concerning the delimitation of its frontiers
or boundaries. In recent years geopolitical entities have become more fluid and this is not reflected consistently
in the tables which were completed at different times and use data of different ages.
A catalogue record for this book is available from the British Library
Contents
How to Use this Book
World Conservation Monitoring Centre
Acknowledgements
Preface
Biodiversity - An Overview
Part 1. BIOLOGICAL DIVERSITY
SYSTEMATICS AND DIVERSITY
he
GENETIC DIVERSITY
The nature and origin of genetic variation
Measuring genetic variation
The interpretation of variation
The environment and the distribution of genetic variation
Conclusion
SYSTEMATICS AND DIVERSITY
Biological classification
Phylogenetic relationships and their estimation
From hierarchy to classification
Taxonomic nomenclature and its regulation
Major features of the hierarchy of life
Systematics and the measurement of biodiversity
SPECIES CONCEPTS
What is a species?
Early species concepts
Evolutionary theory and polytypic species
The biological species concept
The phylogenetic species concept
Species in practice
Conclusion
SPECIES INVENTORY
Current status
Prediction from the existing partial inventory
Other approaches to predicting patterns
Uncharted realms of species richness
Sampling the hyper-diverse but poorly known
New data on tropical insects and what they convey
Prospects for improved species richness estimates
SPECIES DIVERSITY
ae
SPECIES DIVERSITY: AN INTRODUCTION
A brief history of diversity
Measuring biological diversity
The global distribution of species richness
Species and energ
Explanations and Henatieses
iii
Viii
DAWWR
13
17
10.
11.
12.
13.
14.
MICROORGANISMS
Taxonomic scope
Assessment of diversity
Species concepts in microorganisms
Extent of genetic diversity
Regions and habitats of maximum diversity
Role of microorganisms in biodiversity maintenance
Role of microorganisms in biosphere functions
Potential contribution of microorganisms to sustainable development
The need for diversity amongst microorganisms
Ex situ conservation of microorganisms
The taxonomic challenge
LOWER PLANT DIVERSITY
Bryophytes
Lichens
Algae
HIGHER PLANT DIVERSITY
The groups of higher plants
The distribution of higher plants
NEMATODES
Nematode diversity
The ecological importance of nematodes
DEEP-SEA INVERTEBRATES
Deep-sea communities
Ocean trenches
Hydrothermal vents
Cold seeps
SOIL MACROFAUNA
Soil and soil fauna
Patterns of species richness
General patterns of diversity
FISHES
The diversity of fishes
Freshwater fishes: species richness and endemism
Subterranean fishes
Coral reef fishes
HIGHER VERTEBRATES
The groups of higher vertebrates
The distribution of higher vertebrates
ISLAND SPECIES
Plants on oceanic islands
Land snails
CENTRES OF SPECIES DIVERSITY
Introduction
Methods of determining areas of conservation priority
Centres of plant diversity
Centres of avian endemism
SPECIES LOSS
16.
17.
SPECIES EXTINCTION
How species become vulnerable to extinction
A brief history of extinctions
Extinctions in recent history
Current and future extinction rates
THREATENED SPECIES
What is a threatened species?
Globally threatened animals
Aquatic habitats
Threatened species on islands: plants
Threatened species on islands: birds
HABITATS AND ECOSYSTEMS
18.
19.
20.
21.
22:
23.
24.
GLOBAL HABITAT CLASSIFICATION
Ecosystem mapping
Estimating rates of change of ecosystems
BIODIVERSITY AND GLOBAL CLIMATE CHANGE
Modelling global climate change
Effects of climate change on vegetation zones and biodiversity
TROPICAL MOIST FORESTS
What are tropical moist forests?
The global area of tropical moist forest
Factors involved in changes in forest cover
Management practices in tropical forests
Case studies
GRASSLANDS
The world area of grassland
Origins and floral diversity of grasslands
The 20th century impact on grasslands
Case studies
WETLANDS ,
Global extent and distribution of wetlands
Regional extent and distribution of wetlands
Values and threats
Loss of wetlands
Conservation of wetlands
CORAL REEFS
Occurrence of reefs
Values and threats
MANGROVES
The mangrove habitat
Value of mangroves :
Threats to mangrove habitats
25. PLANT USE 331
Food plants 331
Timber 342
Rattans 350
Medicinal plants 350
Ornamental plants 353
26. ANIMAL USE 359
Introduction 359
Food: terrestrial animals 359
Food: fisheries 365
Non-food uses 374
Domestic livestock 389
VALUING BIODIVERSITY
27: BIODIVERSITY AND ECONOMICS 407
Valuing the environment 407
Loss of biodiversity as an economic process 409
Current uses of diverse resources 411
Community use of wildlife resources 411
Ecotourism 413
Existence values 415
The valuation of diverse ecosystems 417
The value of tropical forests 417
The value of wetlands 421
Preserving future options 425
The value of diversity in providing insurance: crop yields 426
Sources of yield variability 429
Crop insurance: the response to increased agricultural risk 430
The value of agricultural genetic diversity 432
The value of biodiversity in the production of pharmaceuticals 434
28. NATIONAL LEGISLATION 441
The protection of wild flora 441
The protection of wild fauna 442
Limitations of species legislation 444
The protection of natural habitats 444
29. PROTECTED AREAS 447
National protected area systems 447
International protected area systems 459
30. MULTILATERAL TREATIES 479
Multilateral treaties 479
vi
31. INTERNATIONAL POLICY AND LEGAL ASSISTANCE
Funding
International obligations: protected areas
Intellectual property rights for biotechnology
Regulated trading in wildlife products
Regional seas programme
32 INTERNATIONAL AID
International development assistance
Bilateral development assistance
Multilateral development assistance
International assistance in forest management
The tropical forestry action plan (TFAP)
The international tropical timber agreement (ITTA)
Debt purchase
33. MANAGEMENT OF INTERNATIONAL RESOURCES
International fisheries management commissions
Antarctica: the evolution of an international resource management regime
34. CURRENT PRACTICES IN CONSERVATION
In situ conservation of threatened plant species
In situ conservation of crops and wild relatives of crops
Institutions involved in ex situ conservation of plants
Techniques for ex situ plant conservation
In situ conservation of animals
Ex situ conservation of animals
Ex situ conservation of animal genetic resources
Ex situ conservation of microbial diversity
352 THE CONVENTION ON BIOLOGICAL DIVERSITY
Background
The biodiversity convention
The biodiversity country studies and unmet financial needs
Future data needs: networking and global monitoring
GLOSSARY
Vii
How to Use this Book
An extensive review of global biodiversity obviously generates substantial quantities of data with the
concomitant problem of how best to present this mass of material. Global Biodiversity is intended to be a
source-book of information and analysis rather than be read cover to cover, so assisting the reader find his/her
way around the book is essential.
The primary means of accessing this wealth of information is through the Contents list (page iii). This is
therefore very detailed and serves some of the function of an index (which it has not been practical to include
because of excess length). The reader is urged to browse the Contents before dipping into the text.
The book is divided into three Parts, each of which opens with a brief overview of its structure and contents.
The Parts are then divided into ten Sections that group together Chapters that address acommon theme. This
structure is outlined below as a guide to the overall organisation of the book.
Part 1. Biological Diversity
e Systematics and diversity
e Species diversity
e Species loss
e Habitats and ecosystems
Part 2. Uses and Values of Biodiversity
e Uses of biological resources
e Valuing biodiversity
Part 3. Conservation and Management of Biodiversity
National policies and instruments
International policies and instruments
Current practices in conservation
Biodiversity Convention
Individual Chapters are divided thematically by major sub-headings, and these are fully listed in the Contents,
which is therefore the key entry point for access to the information.
As far as possible, plain English has been used rather than scientific terminology, but when the use of obscure
technical language has been unavoidable a definition has been provided in the Glossary.
vill
World Conservation Monitoring Centre
The Earth's biological diversity and other natural resources provide many economic, social and aesthetic
benefits to mankind. Effective programmes for sustainable human development must, therefore, incorporate
conservation objectives. Responsible institutions and individuals need access to a service that provides factual
information on conservation issues in a timely, focused and professional way.
This service is provided by the World Conservation Monitoring Centre, Cambridge, UK. Established in
1988 as a company limited by guarantee with charitable status, WCMC is managed as a joint-venture between
the three partners in the World Conservation Strategy and its successor Caring For The Earth: 1\UCN - The
World Conservation Union, UNEP - United Nations Environment Programme, and WWF - World Wide
Fund for Nature. Its mission is to provide information on the status, security, management and utilisation of
the world's biological diversity to support conservation and sustainable development.
To implement this mission, WCMC maintains substantial databases on the status and distribution of plant and
animal species of conservation and development interest; habitats of conservation concern, particularly tropical
forests, coral reefs and wetlands; the global network of national parks and protected areas; and the international
trade in wildlife species and their derivative products. Much of this information is managed with Geographic
Information Systems, and is supported by an extensive bibliography of published and "grey" literature. WCMC
is also involved in providing support for the expansion of national data management and monitoring capabilities
in developing countries, and in developing communication networks for the flow of information.
WCMC contributes its biodiversity data to GEMS - the Global Environment Monitoring System, co-ordinated
by UNEP. GEMS is a collective programme of the world community to acquire, through global monitoring,
and assessment, the data that are needed for the rational management of the environment. GEMS is an element
of the United Nations Earthwatch programme.
WCMC Biodiversity Report Team
John McComb Project Manager
Dr Brian Groombridge Editor and Research Co-ordinator
Esther Byford Production Supervisor
Crawford Allan Research
John Howland Research
Dr Chris Magin Research
Helen Smith Research
Veronica Greenwood Production
Lindsay Simpson Production
Consultant Assistant Editors
Martin Jenkins (general)
Timothy M. Swanson (economics and policy)
Hugh Synge (plants)
WCMC Staff who contributed to the compilation and review process:
Mike Adam, Clare Billington, Simon Blyth, Gillian Bunting, John Caldwell, Lorraine Collins, Dr Mark
Collins, Mary Cordiner, Helen Corrigan, Robert Cubey, John Easy, Jeremy Harrison, Tim Inskipp, Dr
Timothy Johnson, Beverley Lewis, Dr Richard Luxmoore, Lesley McGuffog, Sheila Millar, Dr Ronald I.
Miller, James R. Paine, Dr Robin Pellew, Corinna Ravilious, Jonathan Rhind, Sarah Skinner, Jo Taylor,
Dr Kerry Walter.
Acknowledgements
The production of this Report has been based largely upon the substantial biodiversity databases that WCMC
manages, supplemented by a major world-wide data gathering and standardisation programme. This
information is drawn from an extensive network of scientists, research workers, park managers, wildlife
authorities, conservation bodies and government organisations. WCMC would like to express its thanks for
the contributions of these individuals and agencies, too numerous to mention individually, without whose
support we would not be able to operate.
WCMC particularly recognises with gratitude the fundamental contribution of both the IUCN Species Survival
Commission (SSC) and the Commission of National Parks and Protected Areas (CNPPA), whose commitment
to WCMC over the years has enabled the Centre to expand its databases. Their data have been used extensively
in the Report. WCMC also acknowledges the support of the IUCN Environmental Law Centre, whose legal
data and expertise have contributed significantly.
In compiling the Report itself, WCMC particularly acknowledges the major contributions of the following
people whose names appear at the end of the relevant chapter (unattributed chapters were provided by WCMC
staff):
Dr John Akeroyd, Bruce Aylward, Dr Keith Banister, Dr Gordon Brent Ingram, Dr B.N.K. Davis, Victoria
Drake, Alan Eddy, B.C. Eversham, Alix Flavelle, Shirra Freedman, D.J. Galloway, Sarah Gammage,
Dr Stephen J.G. Hall, Dr P.N. Halpin, Peter Hammond, David Hanrahan, Dr Caroline Harcourt, Prof. D.L.
Hawksworth, Richard J. Hornby, Nigel Howard, Martin Jenkins, David M. John, Sam Johnston, A.S. Jolliffe,
E.A. Leadlay, Dr Rik Leemans, Mike Maunder, Sara Oldfield, Greg Rose, Timothy M. Swanson, Hugh
Synge, Richard Thomas, Ian Tittley, Susan M. Wells, Dr P.S. Wyse Jackson, Dr R.I. Vane-Wright.
The first four chapters were contributed by staff of The Natural History Museum, London. WCMC is
especially grateful for their assistance, and for the efforts of John Peake (Associate Director, Scientific
Development) in facilitating this collaboration.
In addition, WCMC would like to express thanks to the following who have contributed to the compilation
of this Report in a variety of ways:
Dr Dennis Adams, Suraya Affiff, Dr J.Crinan Alexander, M. Altieri, Dr Chris Andrews, Dr Martin Angel,
Prof. Peter Ashton, Margerita Astralaga. Dr Paul Bamps, Dr John Beard, Dr S. Beck, Dr Henk Beentje,
Dr Colin J. Bibby, Dr Mike Bingham, Dr William Block, Dr Robert Boden, Dr A. Bogan, Dr Attila Borhidi,
Dr Philippe Bouchet, Prof. Loutfy Boulos, Dr David Bramwell, Dr F.J. Breteler, Dr Harold Brookfield,
Dr Dick Brummitt, David Brunner, Dr Francoise Burhenne-Guilmin, Victor Bullen, Dr Yvonne A. Byron,
Dr J. Cardiel, Dr Jan Cerovsky, Jim Chapman, Dr Arthur D. Chapman, Dr A. Cleef, P. Colman, Dr R.
Cowie, Dr Quentin Cronk, Mike J. Crosby, James Culverwell II, Michael Dadd, Dr Patricia Davila, Stephen
D. Davis, D.G. Debouck, Dr Jean-Jacques de Granville, Dr Robert DeFilipps, A. Delsaerdt, Nelly Diego,
Dr C. Dodson, Dr John Dransfield, Dr J. Duivenvoorden, M. Dulude, Dr K. Emberton, Lynne Farrell, Prof.
Philip M. Fearnside, Dr Richard Felger, Rosa M. Fonseca, Dr F.R. Fosberg, Dr F. Friedmann, Dr Ib Friis,
Dr John D. Gage, Dr F. Galena, Dr Rodrigo Gamez, Dr Sam M. Gan III, Dr Martin Gardner, N. Gardner,
Dr Steve Gartlan, Dr Alwin Gentry, Dr David Given, Prof. César G6mez Campo, Dr Roger Good, Dr R.
Gopalan, Dr Frederick Grassle, Peter Green, O. Griffiths, Liz Guerin, Prof. Nimal Gunatilleke, Dr M.
Hadfield, Dr S. Halloy, Dr Ole Hamann, Dr Alan Hamilton, Dr Stephen Harris, Melanie F. Heath, Dr A.N.
Henry, Dr Derral Herbst, Prof. Vernon Heywood, Craig Hilton-Taylor, Dr A. Hoffmann, Dr Martin
W. Holdgate, E. Hoyt, Dr Otto Huber, Prof. Dr Gordon Brent Ingram, Dr Frank Ingwersen, Dr Walter
Ivantsoff, Prof. K. Iwatsuki, Dr Peter Wyse Jackson, Dr J. Jérémie, Prof. Robert Johns, Dr Marshall
Johnston, Dr M. Jorgensen, Dr Calestous Juma, Prof. Horng Jye-Su, Dr Ruth Kiew, Dr T. Killeen, Prof.
V. Krassilov, Dr John Lambshead, Prof. Elias Landolt, Dr R. Lara, Dr John Leigh, Dr David Lellinger,
Christine Leon, Blanca Leén, Dr E. Lleras, Dr Paul V. Loiselle, Adrian J. Long, Francisco Lorea,
Dr Rosemary Lowe-MacConnell, Lucio Lozado, Prof. Grenville Lucas, Olga Herrera-MacBryde, Dr Kathy
MacKinnon, Jane MacKnight, Lynne Maclennan, Dr Domingo Madulid, Mike Maunder, Dr Niall McCarten,
Dr Bill McDonald, Bob McDowall, Jeffrey A. McNeely, Dr Tim Messick, Robert Mill, Dr Kenton R.
Miller, Dr Tony Miller, Danya Miskov, Lino Monroy, P. Mooney, Dr Norman Moore, Prof. P. Morat,
Dr P.B. Mordan, Dr Scott Mori, Dr Larry Morse, Michael Moser, Fred Naggs, Dr David Neill, Dr B.
Nelson, Dr Dan Nicholson, Dr Hans Nooteboom, Dr Rosa Ortiz, Dr Maria Tereza Jorge Padua,
Dr Christopher Page, Dr W. Palacios, Dr Mark Perry, Prof. Ghillean T. Prance, Dr M. Prashanth,
Robert Prescott-Allen, Han Qunli, Dr L. Ramella, Dr Orlando Rangel, Dr Peter Raven, Dr Tony Rebelo,
Marcia Ricci, J. Robertson-Vernhes, Joyce Rushton, Dr B.D. Sharma, Samar Singh, Dr D.K. Singh,
Dr Mark Skinner, Joel Smith, D. Smits, Dr Sy Sohmer, C. Sperling, Alison Stattersfield, Dr George
Staples, Dr G. Stephens, Wendy Strahm, Dr Tod Stuessy, Prof. Dr H. Sukopp, Dr R.W. Sussman, Glen
Swindlehurst, Lesley Taylor, Dr Simon Thirgood, Dr Duncan Thomas, Dr F. Thompson, Dr Jim Thorsell,
Dr Mats Thulin, Dr S. Tillier, Simon Tonge, Dr Shigeru Tsuda, Dr Verena Tunnicliffe, Dr C. Ulloa,
Dr E. Vajravelu, Dr Vu Van Dung, Dr Leo Vanhecke, Jane Villa-Lobos, Dr C. Villamil, Dr J.-F. Villiers,
Dr David Wagner, Dr Warren H. Wagner, Dr H. Waldén. Richard Warner, Dr Tom Wendt, Julie S.
Wenslow, Dr Dagmar Werner, Dr Gerry Werren, Dr Tony Whitten, J.T. Williams, Julia Willison, Dr David
S. Woodruff, Dr Richard Wunderlin, Prof. Wang Xianpu, Dr K.R. Young, Prof. Yang Zhouhuai,
The authors are grateful for the assistance provided by the librarians of the Monks Wood Experimental Station,
the University of Cambridge, the Natural History Museum (General, Zoological and Entomological) and the
British Antarctic Survey.
Finally, WCMC recognises with gratitude the substantial financial contributions made by our sponsors listed
on the title page. Their confidence in our ability to complete such an ambitious project is appreciated. In
particular, WCMC thanks the Overseas Development Administration, UK and especially David Turner, Ian
Haines and Mark Lowcock, together with the Ministry of Foreign Affairs, The Netherlands, particularly Ton
van der Zon and Egbert Pelinck. The Ministry of the Environment, Denmark, through the endorsement of
Veit Koester, also contributed, whilst the World Bank, through Mohan Munasinghe has distributed copies
into the developing world. WCMC expresses its sincere gratitude these organisations and individuals.
Preface
We Need Your Data for Future Editions of this Report
In your hands you now hold the most comprehensive review of global biodiversity ever compiled. It represents
the product of numerous scientists, consultants and research institutes each of whom has generously contributed
data or assistance to the compilation of this Report, together with the substantial information holdings that
WCMC already manages. Yet so vast and diverse are the Earth's living resources - the genes, species and
ecosystems that comprise the planet's biotic wealth - and the threats that these resources now face, that this
massive effort has barely scratched the surface.
To build the information store on which this Report is based, we need your help. WCMC will continue to
expand its global biodiversity database and intends to republish the Report every two years. This volume is
therefore the first of a proposed series that will document changes to the status, utilisation and management
of the world's biological resources. We need your contribution to fuel this expansion. We are embarked
upon a long-term process, the aim of which is to mobilize the substantial amounts of data available throughout
the world to encourage a more enlightened conservation practice. Your piece of the jigsaw puzzle may fit
into the overall picture we are trying to create. If you are able to contribute data to expand this Report, we
urgently want to hear from you - don't quibble with its deficiencies which inevitably are numerous; instead
be more constructive by contributing your specialist knowledge to this global conservation effort. We plan to
distribute the database itself later this year in machine-readable format, and it is not too late to include your
information.
The need for reliable quantitative information about the impact of people upon nature has never been greater.
Good intelligence is the key to good decisions, whether about priorities, policies or investments. We need
to develop data gathering and monitoring capabilities at the local and country levels, particularly in the
developing world, and to build networks for the early-warning of new threats to biodiversity. The realisation
of these needs is encompassed in Agenda 21 of the UN Conference on Environment and Development, in the
Biodiversity Convention, and in the Global Biodiversity Strategy, but the basic common factor for the
implementation of all these initiatives is good information. WCMC will make available its information to
support these global enterprises, but to be really effective, we need your data and your participation.
This process of expanding the global database through the networking of national centres must be linked
directly into the Biodiversity Convention. Despite the delays and frustrations in its negotiation, which are
discussed in Chapter 35, the Convention could provide a potent mechanism for implementing global
conservation and sustainable use of biodiversity. Assuming a Convention is eventually agreed, its
effectiveness will depend upon its access to reliable up-to-date scientific information. WCMC will mobilize
its substantial data holdings to support the Convention: information will be its life-blood and WCMC will act
as the catalyst for its operation by providing a massive blood transfusion.
Robin Pellew World Conservation Monitoring Centre
Director 219 Huntingdon Road
Cambridge
24 April 1992 CB3 ODL
: UK
BIODIVERSITY: AN OVERVIEW
This introduction is intended to map out in general terms
some of the principal themes to be encountered in the field
of biological diversity. It will provide a context for the
remainder of the report, in which many of these themes
are further developed.
WHAT IS BIODIVERSITY?
The word ~ biodiversity’ is a contraction of biological
diversity. Diversity is a concept which refers to the range
of variation or differences among some set of entities;
biological diversity thus refers to variety within the living
world. The term ~ biodiversity’ is indeed commonly used
to describe the number, variety and variability of living
organisms. This very broad usage, embracing many dif-
ferent parameters, is essentially a synonym of ~ Life on
Earth’.
Management requires measurement, and measures of diver-
sity only become possible when some quantitative value can
be ascribed to them and these values can be compared. It is
thus necessary to try and disentangle some of the separate
elements of which biodiversity is composed.
It has become a widespread practice to define biodiversity
in terms of genes, species and ecosystems, corresponding
to three fundamental and hierarchically-related levels of
biological organisation.
Genetic diversity
This represents the heritable variation within and between
populations of organisms. Ultimately, this resides in vari-
ations in the sequence of the four base-pairs which, as
components of nucleic acids, constitute the genetic code.
New genetic variation arises in individuals by gene and
chromosome mutations, and in organisms with sexual
reproduction can be spread through the population by
recombination. It has been estimated that in humans and
fruit flies alike, the number of possible combinations of
different forms of each gene sequence exceeds the number
of atoms in the universe. Other kinds of genetic diversity
can be identified at all levels of organisation, including
the amount of DNA per cell, and chromosome structure
and number.
This pool of genetic variation present within an inter-
breeding population is acted upon by selection. Differen-
tial survival results in changes of the frequency of genes
within this pool, and this is equivalent to population
evolution. The significance of genetic variation is thus
clear: it enables both natural evolutionary change and
artificial selective breeding to occur.
Only a small fraction (often less than 1%) of the genetic
material of higher organisms is outwardly expressed in
the form and function of the organism; the purpose of the
remaining DNA and the significance of any variation
within it is unclear.
Each of the estimated 10° different genes distributed
across the world's biota does not make an identical
contribution to overall genetic diversity. In particular,
those genes which control fundamental biochemical proc-
esses are strongly conserved across different taxa and
generally show little variation, although such variation
that does exist may exert a strong effect on the viability
of the organism; the converse is true of other genes.
Further, an astonishing amount of molecular variation in
the mammalian immune system, for example, is possible
on the basis of a small number of inherited genes.
Species diversity
Perhaps because the living world is most widely consid-
ered in terms of species, biodiversity is very commonly
used as a synonym of species diversity, in particular of
“species richness', which is the number of species in a
site or habitat. Discussion of global biodiversity is typi-
cally presented in terms of global numbers of species in
different taxonomic groups. An estimated 1.7 million
species have been described to date; estimates for the total
number of species existing on earth at present vary from
five million to nearly 100 million. A conservative working
estimate suggests there might be around 12.5 million. In
terms of species number alone, life on earth appears to
consist essentially of insects and microorganisms.
The species level is generally regarded as the most natural
one at which to consider whole-organism diversity. Spe-
cies are also the primary focus of evolutionary mecha-
nisms, and the origination and extinction of species are
the principal agents in governing biological diversity in
most senses in which the latter can be defined. On the
other hand, species cannot be recognised and enumerated
by systematists with total precision, and the concept of
what a species is differs considerably between groups of
organisms.
Further, a straightforward count of the number of species
only provides a partial indication of biological diversity,
for implicit within the term is the concept of degree or
extent of variation; that is, organisms which differ widely
from each other in some respect by definition contribute
more to overall diversity than those which are very
similar.
The more different a species is from any other species (as
indicated, for example, by an isolated position within the
taxonomic hierarchy), then the greater its contribution to
any overall measure of global biological diversity. Thus,
the two species of Tuatara (genus Sphenodon) in New
Zealand, which are the only extant members of the reptile
order Rhynchocephalia, are more important in this sense
than members of some highly speciose family of lizards.
Developing this argument, a site with many different
higher taxa present can be said to possess more ftaxo-
nomic diversity than another with fewer higher taxa but
many more species. Marine habitats frequently have
more different phyla but fewer species than terrestrial
habitats; i.e. higher taxonomic diversity but lower
species diversity. Measures under development endeav-
our to incorporate quantification of the evolutionary
uniqueness of species.
The ecological importance of a species can have a direct
effect on community structure, and thus on overall bio-
logical diversity. For example, a species of tropical rain
forest tree which supports an endemic invertebrate fauna
of a hundred species evidently makes a greater contribu-
tion to the maintenance of global biological diversity than
a European alpine plant which may have no other species
wholly dependent on it.
Ecosystem diversity
The quantitative assessment of diversity at the ecosystem,
habitat or community level remains problematic. Whilst
it is possible to define what is in principle meant by genetic
and species diversity, and to produce various measures
thereof, there is no unique definition and classification of
ecosystems at the global level, and it is thus difficult in
practice to assess ecosystem diversity other than on a local
or regional basis and then only largely in terms of vege-
tation. Ecosystems further differ from genes and species
in that they explicitly include abiotic components, being
partly determined by soil parent material and climate.
Ecosystem diversity is often evaluated through measures
of the diversity of the component species.. This may
involve assessment of the relative abundance of different
species as well as consideration of the types of species. In
the first instance, the more equally abundant different
species are, then in general the more diverse that area or
habitat is considered to be. In the second instance, weight
is given to the numbers of species in different size classes,
at different trophic levels, or in different taxonomic
groups. Thus a hypothetical ecosystem which consisted
only of several species of plants, would be less diverse
than one with the same number of species but which
included animal herbivores and predators. As different
weightings can be given to these different factors when
estimating the diversity of particular areas, there is no one
authoritative index for measuring diversity. This obvi-
ously has important implications for the ranking of differ-
ent areas.
Biodiversity: its meaning and measurement
The differences between these conceptual perspectives on
the meaning of biodiversity, and the associated semantic
problems, are not trivial. Management intended to main-
tain one facet of biodiversity will not necessarily maintain
another. For example, a timber extraction programme
which is designed to conserve biodiversity in the sense of
site species richness may well reduce biodiversity meas-
ured as genetic variation within the tree species harvested.
Clearly, the maintenance of different facets of biodiversity
will require different management strategies and re-
sources, and will meet different human needs.
Even if complete knowledge of particular areas could be
assumed, and standard definitions of diversity be derived,
the ranking of such areas in terms of their importance with
respect to biological diversity remains problematic. Much
depends on the scale that is being used. Thus, the question
of what contribution a given area makes to global biologi-
cal diversity is very different from the question of what
contribution it makes to local, national or regional bio-
logical diversity. This is because, even using a relatively
simplified measure, any given area contributes to biologi-
cal diversity in at least two different ways - through its
richness in numbers of species and through the endemism
(or geographical uniqueness) of these species. The relative
importance of these two factors will inevitably change at
different geographical scales, and sites of high regional
importance may have little significance at a global level.
Neither of these factors include any explicit assessment of
genetic diversity.
Although the word biodiversity has already gained wide
currency in the absence of a clear and unique meaning,
greater precision will be required of its users in order that
policy and programmes can be more efficiently defined in
the future.
BIODIVERSITY: CHANGES IN TIME AND
SPACE
Changes over time
The fossil record of life in geological time is very incom-
plete. There is marked variation between higher taxa and
between species in different ecosystems in the extent to
which individuals are susceptible to preservation and to
subsequent discovery. Chance factors have played a large
part, and interpretation by palaeontologists of the avail-
able material is beset by differences of opinion. Thus, the
record is relatively good for shallow-water hard-bodied
marine invertebrates, but poor for most other groups, such
as plants in moist tropical uplands.
Two salient points appear well-substantiated. Firstly,
taxonomic diversity, as measured by the number of rec-
ognised phyla of organisms, was greater in Cambrian
times than in any later period. Secondly, and keeping in
mind the difficulty of disentangling artifacts of the record
from the underlying pattern, it appears that species diversity
and number of families have undergone a net increase
between the Cambrian and the Pleistocene epoch, although
interrupted by isolated phases of mass extinction (few of
which are reflected in the fossil record of plants).
Changes in space
In general, species diversity in natural habitats is high in
warm areas and decreases with increasing latitude and
altitude. On land, diversity is also usually higher in areas
of high rainfall and lower in drier areas. The richest areas
are undoubtedly tropical moist forests. If current estimates
of the number of species (mainly insects) comprising the
microfauna of tropical moist forests are credible, then
these areas, which cover perhaps 7% of the world's
surface area, may well contain over 90% of all species.
If the diversity of larger organisms only is considered,
then coral reefs and, for plants at least, areas with Medi-
terranean climate in South Africa and Western Australia,
may be as diverse. Gross genetic diversity and ecosystem
diversity will, by definition, tend to be positively corre-
lated with species diversity (although there are indications
that some tropical species show more genetic diversity
than related temperate species, and some habitat general-
ists more than habitat specialists).
The reasons for the large-scale geographic variation in
species diversity, and in particular for the very high
species diversity of tropical moist forests, are not fully
understood and involve two interconnected questions: the
origin of diversity through the evolution of species and
the maintenance of diversity. Both these involve consid-
eration of the present and historic (in a geological or
evolutionary sense) conditions prevailing in particular
areas, principally climatic but also edaphic and topo-
graphic. Climatically benign conditions (warmth, mois-
ture and relative aseasonality) over long periods of time
appear to be particularly important.
It is often assumed that areas with so-called climax
ecosystems will be more diverse than areas at earlier
successional stages. However, an area with a mosaic of
systems at different successional stages will probably be
more diverse than the same area at climax provided that
each system occupies a sufficiently large area of its own.
In many instances, human activities artificially maintain
ecosystems at lower successional stages. In areas that have
been under human influence for extended periods, notably
in temperate regions, maintenance of existing levels of
diversity may involve the maintenance of at least partially
man-made landscapes and ecosystems, mixed with ade-
quately sized areas of natural climax ecosystems.
Loss of biodiversity
The loss of biological diversity may take many forms but
at its most fundamental and irreversible it involves the
extinction of species.
Over geological time, all species have a finite span of
existence. Species extinction is therefore a natural process
which occurs without the intervention of man. However,
it is beyond question that extinctions caused directly or
indirectly by man are occurring at a rate which far exceeds
any reasonable estimates of background extinction rates,
and which, to the extent that it is correlated with habitat
perturbation, must be increasing.
Unfortunately, quantifying rates of species extinction,
both at present and historically, is difficult and predicting
future rates with precision is impossible.
Documenting definite species extinctions is only realistic
under a relatively limited set of circumstances, where a
described species is readily visible and has a well-defined
range which can be surveyed repeatedly. Unsurprisingly,
most documented extinctions are of species that are easy
XV
to record (e.g. land snails, birds) and inhabit sites which
can be relatively easily inventoried (e.g. oceanic islands).
The large number of extinct species on oceanic islands is
not solely an artefact of recording, because island species
are generally more prone to extinction as a result of human
actions.
Rather than being derived from observed extinctions,
therefore, quoted global extinction rates are derived from
extrapolations of measured and predicted rates of habitat
loss, and estimates of species richness in different habi-
tats. These two estimates are interpreted in the light of a
principle derived from island biogeography which states
that the size of an area and of its species complement tend
to have a predictable relationship; fewer species are able
to persist in a number of small habitat fragments than in
the original unfragmented habitat, and this can result in
the extinction of species.
Even on best available present knowledge, these estimates
involve large degrees of uncertainty, and predictions of
current and future extinction rates should be interpreted
with very considerable caution. Pursuit of increased ac-
curacy in the estimation of global extinction rates, how-
ever, whilst of great concern, is not a crucial activity; it
is more important to recognise in general terms the extent
to which populations and species which are not monitored
are likely to be subject to fragmentation and extinction.
Loss of biodiversity in the form of crop varieties and
livestock breeds is of near zero significance in terms of
overall global diversity, but genetic erosion in these
populations is of particular human concern in so far as it
has implications for food supply and the sustainability of
locally-adapted agricultural practices. For domesticated
populations, loss of wild relatives of crop or timber plants
is of special concern for the same reason. These genetic
resources may not only underlie the productivity of local
agricultural systems but also, when incorporated in breed-
ing programmes, provide the foundation of traits (disease
resistance, nutritional value, hardiness, etc.) of global
importance in intensive systems and which will assume
even greater importance in the context of future climate
change.
Erosion of diversity in crop gene pools is difficult to
demonstrate quantitatively, but tends to be indirectly
assessed in terms of the increasing proportion of world
cropland planted to high yielding, but genetically uniform,
varieties.
The causes of loss of biological diversity
Species may be exterminated by man through a series of
effects and agencies. These may be divided into two broad
categories: direct (hunting, collection and persecution),
and indirect (habitat destruction and modification).
Overhunting is perhaps the most obvious direct cause of
extinction in animals, as it has affected several large and
well-known species. In terms of overall loss of biodiver-
sity, however, it is undoubtedly far less important than
the indirect causes of habitat modification and loss. Nev-
ertheless, as it self-evidently selectively affects species
which are or have been considered a harvestable resource,
it has important implications for the management of
natural resources.
Genetic diversity, as represented by genetic differences
between discrete populations within wild species, is liable
to reduction as a result of the same factors affecting
species. The genetic diversity represented by populations
of crop plants or livestock is liable to reduction as a result
of mass production; the desired economies of scale de-
mand high levels of uniformity.
Virtually any form of sustained human activity results in
some modification of the natural environment. This modi-
fication will affect the relative abundance of species and
in extreme cases may lead to extinction. This may result
from the habitat being made unsuitable for the species (for
example, clear-felling of forests or severe pollution of
rivers), or through the habitat becoming fragmented. The
latter has the effect of dividing previously contiguous
populations of species into small sub-populations. If these
are sufficiently small, then chance processes lead to raised
probabilities of extinction within a relatively short time.
A major, though at present largely unpredictable, change
in natural environments is likely to occur within the next
century as a result of large-scale changes in global climate
and weather patterns. There is a high probability that these
will cause greatly elevated extinction rates, although their
exact effects are at present unknown.
MAINTAINING BIOLOGICAL DIVERSITY
The maintenance of biological diversity at all levels is
fundamentally the maintenance of viable populations of
species or identifiable populations. This can be carried out
either on site or off site. Some integrated management
programmes have begun to link these basically dissimilar
approaches.
In situ conservation
The maintenance of a significant proportion of the world's
biological diversity at present only appears feasible by
maintaining organisms in their wild state and within their
existing range. This is generally preferable to other
courses of action because it allows for continuing adapta-
tion of wild populations by natural evolutionary processes
and, in principle, for current utilisation practices to con-
tinue (although these often require enhanced manage-
ment).
Ex situ conservation
Viable populations of many organisms can be maintained
in cultivation or in captivity. Plants may also be main-
tained in seed banks and germplasm collections; similar
techniques are under development for animals (storage of
embryos, eggs, sperm) but are more problematic. In any
event, ex situ conservation is clearly only feasible at
present for a small percentage of organisms. It is ex-
tremely costly in the case of most animals, and while it
would in principle be possible to conserve a very large
XVi
proportion of higher plants ex situ, this would still amount
to a small percentage of the world's organisms. It often
involves a loss of genetic diversity through founder effects
and the high probability of inbreeding.
WHY CONSERVE BIOLOGICAL DIVERSITY ?
This question can be asked from a number of different
perspectives, all conditioned by a variety of cultural and
economic factors. The various answers given, arguing for
the maintenance of biological diversity, have tended to
become increasingly confused. Different goals have dif-
ferent implications for the elements and extent of biologi-
cal diversity that must be maintained. Among these goals
are the following:
¢ the present and potential use of elements of biodiversity
as biological resources
e the maintenance of the biosphere in a state supportive
of human life
e the maintenance of biological diversity per se, in
particular of all presently living species.
Biological diversity as a resource
It is evident that a certain level of biological diversity is
necessary to provide the material basis of human life: at
one level to maintain the biosphere as a functioning system
and, at another, to provide the basic materials for agricul-
ture and other utilitarian needs.
Food
The most important direct use of other species is as food.
Although a relatively large number of plant species,
perhaps a few thousand, have been used as foodstuffs, and
a greater number are believed to be edible, only a small
percentage of these are nutritionally significant on a global
level, and only very few of these have been intensively
managed on a commercial scale. Similarly, very many
animal species are eaten (mostly fishes), but only a very
small percentage are globally of nutritional significance.
A few dozen species, mostly mammals, are managed in
some kind of husbandry system, and a handful of these
are globally significant.
It is clear that successful cultivation of agricultural crops
on a large scale requires a suite of other organisms (chiefly
soil microorganisms and, in a few cases, pollinators) but
these probably amount to a statistically insignificant per-
centage of global biological diversity. Highly productive
agricultural systems also require the virtual absence of
some elements of biological diversity (pest species) from
given sites.
Whilst relatively little diversity is currently used in com-
mercial food production, the very high probability of
global climate change, predicted to result in large-scale
shifts in natural vegetation and in agricultural systems, has
focused attention on the need for conservation of plant
genetic resources in order to maintain crop productivity
under different climatic regimes. This ~ insurance value’
of diversity is also evident in contemporary conditions,
where increased genetic uniformity is correlated with
increased crop yield variation.
Pharmaceuticals
Medicinal drugs derived from natural sources make an
important global contribution to health care. An estimated
80% of people in less-developed countries rely on tradi-
tional medicines for primary health care; this shows no
signs of decline despite availability of western medicine.
Some 120 chemicals extracted in pure form from around
90 species are used in medicines throughout the world.
Many of these cannot be manufactured synthetically: the
cardiac stimulant digitoxin, the most widely used car-
diotonic in western medicine, is extracted direct from
dried Digitalis (foxglove); synthetic vincristine, used to
treat childhood leukaemia is only 20% as efficacious as
the natural product derived from Catharanthus roseus
(Rosy Periwinkle).
As with agriculture, and excluding traditional medicines,
at present only a very small percentage of the world's
biodiversity contributes on a global scale to health care.
Many argue that technological advances within the phar-
maceutical industry, and in particular those involving the
design and manufacture of synthetic drugs, will mean that
this contribution is more likely to fall than rise. However,
natural diversity might be increasingly valued for the
~blueprints' it provides for new synthetic drugs.
Other material values of biological diversity
Many natural or semi-natural ecosystems, some of which
may be of high biological diversity, are of considerable
benefit to man. Examples are:
e the role of forests in watershed regulation and stabili-
sation of soils in erosion-prone areas
e the role of mangroves in coastal zone stabilisation and
as nursery areas for fisheries species
e the role of coral reefs in supporting important subsis-
tence fisheries
e the role of natural ecosystems protected as national
parks in generating income from wildlife tourism.
In general, however, these values are only indirectly
related to biological diversity. That is, a certain level of
species richness is required for these functions but there
is not necessarily a direct correlation between the value
of the ecosystem and its diversity, nor in all cases do a
particular set of species have to be present. Thus, man-
grove ecosystems are generally of far lower diversity than
adjacent lowland terrestrial forests but in resource terms
are likely to be of comparable value. The savannas of east
and southern Africa, which are of great importance in
generating revenues from tourism, are less diverse than
the moist forests in these countries which have far less
potential for tourism.
The precautionary principle
While it is evident that at present a relatively small
proportion of the world's biological diversity is actively
exploited by man, other elements of biological diversity
may be important for different reasons:
e they have values which are unused or unknown at
present but which could enhance the material well-be-
XVil
ing of mankind if these values were discovered and
exploited
e they may become useful or vital at some time in the
future owing to changing circumstance.
These factors support a precautionary line in maintain-
ing biological diversity - that is, actually or potentially
useful resources should not be lost simply because we
do not know about or value them at present. However,
although this precautionary argument has wide applica-
bility it has limited force. It is based on estimates of the
potential value of a given element of biological diversity
which must be balanced against the actual cost of
maintaining it or refraining from destroying it. Thus,
unless a given element is identified as vital, it must have
a finite value and there must therefore come a point at
which the projected costs required to maintain it will
outweigh any probable benefits. The fact that these
costs and benefits are rarely if ever precisely quantifi-
able means that such calculations will involve the esti-
mation of probabilities and risks.
Conclusions on resource values
Experience and general ecological theory indicate that no
single species is indispensable in maintaining basic eco-
logical processes on a global scale and that, in general
terms, the rarer a species is, the less likely it is to play an
important ecological role on even a local level. In other
words, every species has a finite resource value and,
although in some cases this value may be very high, in the
case of increasingly rare species it tends to zero.
Similarly, with respect to species which may be directly
useful to man, chiefly as food and pharmaceuticals, the
vast majority of species can be said with high probability
to have little potential. Experience enables us to identify
those groups of taxa where there is a higher probability
of value (e.g. wild relatives of crop species, and certain
plant families for pharmaceuticals).
General conclusions to be drawn from the above discus-
sion may be that considering species only as material
resources, it would be more cost-effective to:
e maintain systems and areas rich in species than those
poor in species
e maintain those known to be useful, or regarded as
having a high probability of being useful, than to
maintain other species.
These conclusions indicate that resource values of biodi-
versity, and in particular the cost-benefit approach to
conservation, do not of themselves provide justification
for the wide-ranging approach to biodiversity conserva-
tion that many seek to pursue. Such arguments must
have limited applicability and limited force, and consid-
erable caution must be exercised when citing them, es-
pecially when extrapolating from the particular (the
rationale for maintaining particular species or a certain
level of biological diversity) to the general (that all bio-
logical diversity is inherently valuable as a resource
and must therefore be preserved).
Biodiversity and the biosphere
Human activities are affecting the biosphere on a global
scale. It is important in the present context to establish the
extent to which losses in biological diversity may contrib-
ute to these changes in having an impact on man.
One of the most obvious of such global changes is the
perturbation of the carbon cycle, leading to a steady
increase in atmospheric CO? levels. This will probably
have far-reaching, although at present unpredictable, ef-
fects on global climate patterns which may in turn have
serious consequences for human welfare.
A significant part of this is ascribable to industrial proc-
esses, especially the burning of fossil hydrocarbon fuels
for energy generation. However, it is believed that altera-
tion of existing natural or semi-natural ecosystems is also
important. In particular the large-scale destruction of
tropical moist forests is implicated, both in contributing
to atmospheric CO? through burning and in decreasing the
carbon-fixing potential of the biosphere. The high risk of
serious consequences for humans of global climate
changes is itself a strong argument for decreasing rates of
forest clearance. It must, however, be stressed that this
argument applies to tropical moist forest as ~ forest', rather
than as “a highly diverse ecosystem’. Diversity is impor-
tant only to the extent that it contributes to the system
functioning as a carbon sink and the argument applies
equally to other systems with a similarly high capacity for
carbon fixation, such as tropical freshwater swamps,
although these are far less diverse than tropical moist
forest. In more general terms, there appears to be no direct
or obvious link between the importance of an ecosystem
in maintaining essential global ecological processes and
its diversity, although more research is required.
Non-resource values of biological diversity
It is evident that resource-based arguments for the main-
tenance of biological diversity have very considerable but
finite force; therefore any fundamental justification for
striving to maintain all currently existing biological diver-
sity must lie outside the realm of material resource values.
Such justification usually devolves onto two principles -
ethics and aesthetics - which themselves lie outside the
realm of science.
Ethics
For some cultures, ethical beliefs provide the strongest
grounds for maintaining biological diversity, and indeed
in some eastern countries much of the remaining diversity
in densely populated areas can be attributed directly to
religious practices. However, without recourse to an
absolutist moral code, it is difficult to argue compellingly
for an ethical imperative for the maintenance of all
existing biological diversity. Whilst the killing of any
living organism may be morally unacceptable to some
people, there are problems in extending this argument to
the conservation of biological diversity. At an extreme
level, any individual organism that is not genetically
identical to another represents a facet of this diversity, and
XViil
a strict ethical argument would proscribe its destruction.
It may be understandable to object to the killing of an
elephant on moral grounds, but is it any less moral to eat
wheat, which is grown from genetically diverse seeds,
than to eat potatoes, most of which are grown from
genetically identical clones? Similarly, there are difficul-
ties in demonstrating that a species, which is to some
extent a human construct, has any greater “right' to
existence as an entity than any one of the individuals of
which it is comprised.
Neverthess, the fact remains that ethics provides a pow-
erful argument against the destruction of biological diver-
sity. In practice, this argument is often contingent on other
grounds, particularly the precautionary principle. For
example, it may be considered immoral to destroy some-
thing which is now, or may be in the future, regarded as
valuable to others. This is embodied in the ~ stewardship’
argument. The principle of inter-generational responsibil-
ity underpins the ethical case for conservation in the
developed world, although it may be of little practical
relevance to a desperate farmer faced with the reality of
survival in a developing country.
Aesthetics
Arguments for the maintenance of biological diversity for
its aesthetic appeal are compelling but have limited force,
as they must be dependent on relative aesthetic judge-
ments. Such judgements could presumably discard some
organisms (those not visible, for example) as not worthy
of being maintained. They are also unlikely to hold sway
in the face of counter arguments that certainly exist for
the destruction in the wild of harmful organisms, such as
malarial Plasmodium species. Further, because genetic
diversity is not susceptible to aesthetic appreciation, aes-
thetic criteria can be applied only to species and ecosystem
aspects of biodiversity.
Regardless of individual aesthetic judgements, it is un-
doubtedly the case that humans very strongly favour
variety in most areas of their experience. This need is
particularly evident in the realm of the natural world. That
is, diversity itself, and biological diversity in particular,
is held in some poorly-definable but fundamental sense to
be a highly desirable phenomenon. This is no mere notion,
but a need that is very deeply felt, and a fundamental part
of the spiritual life of many people. It is not important that
the reasons for this cannot be fully articulated; the need
is strongly manifest and should have force in determining
action.
Overall, while it is evident that neither ethical nor aes-
thetic arguments provide of themselves sufficient grounds
for attempting to maintain all existing biological diversity,
a more general and pragmatic approach recognises that
different but equally valid arguments (resource values,
precautionary values, ethics and aesthetics, and simple
self-interest) apply in different cases, and between them
provide an overwhelmingly powerful case for biodiversity
conservation.
PART 1
BIOLOGICAL DIVERSITY
Part 1 introduces some of the principal elements comprising biological diversity.
Where appropriate, it discusses the ways in which they are measured, their patterns of
distribution in space and the changes they have undergone over time, and notes their
ecological importance. The main emphasis is on diversity at the species level. The
chapters in Part 1 are grouped into four sections.
The first section (Chapters 1-4) is concerned broadly with the science of systematics
as the primary approach to biodiversity. The opening three chapters cover: genetic
diversity among species and populations, the scope and practice of systematics, and
the meanings of the word ‘species’. Although most debate about biodiversity has been
in terms of species, it is important to recognise that the ‘species’ is not a standard
unit; the way species are defined differs between groups and between taxonomists.
The fourth chapter deals in considerable detail with the complex topic of global
species numbers: how many species have been named and how many species probably
exist but are as yet unknown and undescribed? There is considerable uncertainty about
the number of valid described species, and extreme uncertainty about the global
species total: conservative working estimates suggest 1.7 million described species and
12.5 million in total (estimates of the latter range up to 100 million).
The second section (Chapters 5-15) presents a review of biodiversity at the species
level. Chapter 5 provides a general introduction to the subject of species diversity,
while Chapters 6 to 14 present a series of case studies of different taxonomic or
ecological groups. Many of the data sets presented here are entirely new. No attempt
has been made systematically to cover all organisms in a consistent manner. The
groups included and the kinds of data presented have to a great extent been dictated
by the availability of information and expertise, although we have tried to cover some
groups and communities that are less familiar, or highly diverse, or both. Species
richness of tropical forest insects is discussed at length in Chapter 4, along with an
outline of sampling procedures which could result in much-improved data on their
distribution. Groups that have not received detailed review will be considered in
future editions of this report.
In this section, Chapter 8, on ferns, gymnosperms and flowering plants, and Chapter
13, on vertebrates (excluding fishes), include large data tables which attempt to give
an estimate, for each major group, of the total number of species in each country of
the world, and an estimate of the number endemic (restricted) to each country.
Chapter 12 includes data tables of freshwater fish species number and endemism in
rivers and lakes.
This section closes with a discussion (Chapter 15) of some of the ways in which data
on species distribution can be analysed to identify sites or areas which are particularly
rich in species or contain a high proportion of endemic species. Conservation of these
areas will be particularly important in efforts to maintain global biodiversity. This
approach is illustrated by data derived from two global-level projects dealing with
plants and with birds.
The third section contains two chapters which deal with trends in species diversity
over time. Chapter 16 introduces the phenomenon of extinction; while extinctions in
palaeontological time are discussed, the main emphasis is on historical and recent
extinction, and the problems of predicting current and future rates of species loss. An
attempt to list the animal species known to have become extinct since 1600 is included
in this chapter. Chapter 17 discusses species threatened with extinction, in particular
those which have been assigned to one of the IUCN threatened species categories. It
covers the taxonomic, habitat and geographic distribution of species listed by [UCN as
threatened, and discusses the factors leading to population decline.
The fourth and final section moves on to look at the habitat and ecosystem level of
biodiversity. The opening chapter (18) introduces the theme of global community
classifications, and notes some of the conceptual and practical difficulties which hinder
their construction. Chapter 19 briefly outlines evidence for global climate change, and
its predicted impact on protected areas. Both these chapters are illustrated by full
colour maps.
Chapters 20 to 24 in turn cover five ecosystem types: tropical rain forest, grassland,
wetlands, coral reefs and mangrove forest. A selection of systems which are species-
rich or under particular threat have been included; no attempt has been made
systematically to review all ecosystems (others will be included in future editions).
Chapter 20, on tropical forests, discusses in some detail the various attempts that have
been made to estimate the rate at which this habitat is being modified, and the
difficulties inherent in such estimation. This should be read in conjunction with
Chapter 4, on species inventory, and Chapter 16, which in discussing estimates of
current and future rates of extinction, notes that no precise quantitative link can be
made between species number in tropical forests, rates of forest loss, and rates of
species extinction.
1. GENETIC DIVERSITY
This section introduces concepts from genetics necessary for
an understanding of the generation and maintenance of
biological diversity.
THE NATURE AND ORIGIN OF GENETIC
VARIATION
Genes are the blueprints that make us and all the other
organisms around us what we are. They consist of a
discrete segment of deoxyribonucleic acid (DNA), a linear
molecule composed of sequences of four different
nucleotide bases. From the seemingly simple code contained
in the sequence of these four bases of DNA comes the
overwhelming complexity and diversity of the living world.
Living organisms can be divided very broadly into
eukaryotes, in which the cell nucleus is bounded by a
membrane, contains a number of organelles, and has its
DNA combined with proteins to form chromosomes, and
prokaryotes, in which these features are lacking. All higher
organisms are eukaryotes; bacteria are prokaryotes.
Bacteria generally have a single copy of each of their genes
located on a single piece of DNA and usually they tend to
reproduce asexually, that is without the coming together of
genetic information from another individual. Sometimes
bacteria obtain some or all of the genetic material from
other individuals in a process analogous to sexual
reproduction in animals and plants. Thus, concepts of
species developed principally with reference to higher
organisms do not apply exactly to bacteria. Work is just
beginning to characterise the nature and extent of genetic
variation in a few bacteria. Given the huge diversity that
has evolved over three billion years it is not surprising that
bacteria appear to be a very complex group.
Genes are arranged linearly along the DNA and in most
eukaryote organisms there are something like 50,000 of
them. The actual quantity of DNA in each cell of different
species of eukaryotes varies over three orders of magnitude
(Fig. 1.1). Much of this DNA is not coding for anything
and it is still an active area of research to understand what,
if anything, all this apparently ‘extra’ DNA is doing. Our
ignorance of its function, however, does not stop it from
being useful for answering some kinds of questions, as
discussed below. In most of the organisms we can see with
the naked eye (animals and plants) the DNA of a cell is
divided among a number of chromosomes. Humans have 23
different chromosomes. These chromosomes generally exist
in two copies within each cell of the body and the organism
is then said to be diploid; thus humans have a total of 46
chromosomes per cell. For the majority of organisms,
which have sexual reproduction, one of these copies comes
from the mother and the other from the father. Sex in
genetic terms is just this, the coming together of genetic
information from separate individuals. In this way genetic
differences from different individuals may be combined in
their offspring to produce new combinations upon which
evolutionary processes can work. Asexually reproducing
organisms must wait for the occurrence of different
mutations in the same lineage to achieve these new
combinations of genes.
Genetic Diversity
Mutations are changes in the DNA. They occur in many
ways. Mutations produce variation and variation is the raw
material of evolution. The same gene can exist in a number
of variants and these variants are called alleles. If the two
copies of a particular gene possessed by an individual are
different alleles, the individual is said to be heterozygous at
that gene. If the two copies are the same ailele the
individual is homozygous at that gene. A population of a
species that has more than one allelic form of a particular
gene is said to be polymorphic for that gene. If there are
two alleles for a gene there are two possible homozygotes
and one heterozygote. If there are three alleles, there are
three homozygotes and three heterozygotes. For four alleles
there are four homozygotes and six heterozygotes, and so
on. Now consider the possibilities when we look at two
polymorphic genes, and three, and on to the thousands that
are polymorphic in most outbreeding organisms.
The number of possible combinations is vast - much larger
than the number of individuals making up a species. This is
the variation that the evolutionary process works on, and
that provides the production attributes which agricultural
development seeks to incorporate into crop varieties and
livestock breeds.
The material below considers what is known of the
implications of all this variation, how it changes and
spreads, and the effects of human activities on genetic
diversity and evolutionary processes.
MEASURING GENETIC VARIATION
Measurements of genetic variation are useful for studies of
two broad classes of problems. One of these is the testing
of theories about the nature of the forces acting on genetic’
variants - the nuts and bolts of evolution. There is a large
body of mathematical and statistical theory about the
genetics of populations, the basis of which was formulated
by 1930. Only now, with the advent of DNA technology,
do we have sufficiently powerful tools to begin rigorously
testing these theories and their more recent elaborations.
The other class of problems uses measures of genetic
variation as a tool for understanding relationships among
organisms and the diversity within and divergence between
them.
There are necessarily important connections between the
two sets of problems. Indeed, the central debate in
evolutionary genetics is about whether most of the genetic
variation seen in natural populations is maintained by
natural selection or is neutral and therefore is subject only
to the laws of chance. The issues at stake in this debate are
crucial to the understanding of the mechanisms of the
evolutionary process but they are not so important in the
very practical matters of assessing differences between
individuals, populations and species that are our main
concerns here.
Allozymes
The first widely applicable technique for measuring genetic
variation does so at one remove from the DNA itself. This
Part 1. Biological Diversity
Figure 1.1
PROTISTS
Euglenozoa
Ciliophora
Sarcodina
FUNG!
ANIMALS
Sponges
Anne lids
Molluscs [—
Crustaceans [—
Insects [—
Echinoderms [—
Agnathes [—
Taxon
Sharks/ Rays i=
Bony Fish [-
Amphibians [—
Reptiles [—
Birds |—
Mammals [f—
PLANTS ;—
Algae
Pter idophytes
Gymnosperms [—
Angiosperms |—
Range of DNA content in eukaryote organisms
1 walt rit t {Let ts| rit ve | 1 Ji} ed te | 1 paneer eee
1 10 100 1,000 10, 000 100,000 1000, 000
Range of DNA content wm Range
Source: from data tabulated by Li, W. and Graur, D. 1991. Fundamentals of Molecular Evolution. Sinauer Associates Inc., Sunderland, Mass.
technique is protein electrophoresis and it depends on the
differences in electrical charge between variants of specific
enzymes (allozymes) coded for by DNA. These charge
variants migrate at different rates in gels subjected to an
electric field and can therefore be differentiated from one
another. It was this method that first revealed that on
average 20-30% of the proteins of most organisms exist in
more than one allelic form. This level of variation was not
expected and the search for adequate explanations for it has
been a major force in evolutionary genetics for more than
20 years.
By measuring the frequencies of different variants in groups
of individuals sampled from different areas we can quantify
the amount of variation within and between individuals and
thereby get a picture of the geographic structure of the
species in genetic terms. Not all changes in DNA result in
a charge change which allows variants to be separated on a
gel but this method still provides a good approximation to
changes at the DNA level, at least within species and
between closely related species. At greater taxonomic
distances the probability of two different variants showing
the same mobility in the gel system becomes high enough
for the method to break down. Nevertheless, analysis of
allozyme frequencies still has an important role to play in
the study of intraspecific variation and the bulk of available
data on genetic variation comes from studies of frequencies
of electrophoretic variants of a number of enzymes.
Very recently development of DNA technology has
provided us with the means to sample genetic variation
directly at the DNA level. At present, however, these
methods are more expensive and generally more difficult to
perform than allozyme techniques. Therefore there are not
yet the large amounts of data on within-species variation
available from allozyme studies. This situation is changing
rapidly as the necessary technology becomes more widely
available and less expensive. The following sections give
brief descriptions of the main techniques of use in
phylogenetic and population genetic studies.
Because different parts of the DNA evolve at different rates
we can choose to study particular segments to answer
particular questions. Some genes change very slowly and
can be used to study relationships among groups of
organisms which diverged from one another hundreds or
even thousands of millions of years ago. Other regions of
DNA change at such a rapid rate that every individual in a
population, except for identical twins and other such clones,
is distinct. Still other regions of DNA show intermediate
levels of variability which are useful for studies of variation
within and between populations of a species, or of variation
between closely related species.
Restriction fragment polymorphisms (RFLPs)
The DNA-based techniques most widely used for studies of
within- and between-population variation make use of the
properties of enzymes derived from various species of
bacteria which use them to protect themselves from
infection by viruses by cutting (restricting) invading viral
DNA. These restriction enzymes are very specific in the
DNA sequence they recognise and cut, and they form the
backbone of the technology of DNA manipulation. If DNA
from an individual is extracted and cut with a restriction
enzyme and the resulting fragments separated by length in
an electrophoretic gel a pattern is obtained. Another
individual may have a change in its DNA which produces
an additional site recognised by the enzyme, or it might
have changed in such a way that a recognition site has
disappeared, thereby changing the pattern of restriction
fragments seen on a gel. By repeating this process with
other individuals and restriction enzymes, patterns of
variation can be seen and analyzed to estimate the amount
of variation in the DNA sequences among the individuals.
These restriction fragment length polymorphisms (RFLPs)
are very useful for determining the geographic structure of
populations. By measuring the frequencies of different
patterns in populations of a species we can estimate the
amount of gene flow or genetic cohesion among the
populations.
DNA sequencing and the polymerase chain reaction
Another more powerful (and more expensive) method of
assessing genetic variation is to sequence a portion of the
DNA itself. With the advent of the polymerase chain
reaction technique (PCR), which can be used to make
millions of copies of a particular region of DNA, it is now
possible to obtain DNA sequence data from a wider variety
of organisms much more quickly than was possible
previously. The exquisite sensitivity of the PCR permits the
amplification of a sequence from minute amounts of starting
material - as little as a single cell. This has very important
implications for obtaining data from very small organisms
which contain too little tissue to use with RFLPs, and from
larger organisms without having to kill or otherwise injure
them. A minute drop of blood or a hair root or a feather
are now adequate material for DNA sequence-based work.
This has obvious importance in dealing with rare and
endangered species.
THE INTERPRETATION OF VARIATION
Different measures of variation can be used to investigate
relationships ranging from very distant groups, such as
phyla, to closely related individuals within a population.
Often an understanding of relationships among closely
related individuals is necessary for understanding behaviour
and evolutionary processes within a species. Similarly, with
breeding programmes for endangered species it is important
to know the degree of genetic relatedness of individuals so
that deleterious effects from inbreeding can be minimised.
The technique of genetic fingerprinting can provide this
information. Fingerprinting makes use of a common but
peculiar group of DNA sequences known as minisatellites.
These are dispersed throughout the genome and consist of
tandemly repeated copies of short sequence units. High
levels of variation in the numbers of these repeated units
are exploited in fingerprinting to identify close relatives.
Genetic Diversity
Biologists have long wanted to know if the genetic
differences between species were of a different sort from
the differences between individuals within a species. The
answer appears to be that interspecific differences are not
different in kind from intraspecific variation. Animal
species usually differ at a large number of genes; single
mutations are seldom, if ever, responsible for speciation
events. The genetics of speciation is not discussed here
although information on genetic distance between species-
level populations in selected vertebrate genera, derived from
methods outlined above, is show in Fig. 1.2.
In an outbreeding species every individual has a unique
combination of alleles and the shuffling of genes that occurs
in sexual reproduction insures that every future individual
will be unique as well. If every individual is unique, what
use are genetic data in making decisions about conservation
problems? This question gets us to the heart of some
fundamental problems in biology. Our knowledge of how a
genotype is translated into a phenotype, a body, is very
sketchy and this is an area of major research effort in
biology. Genetic criteria for uniqueness and justification for
conservation are not simple problems. In the sections below
we will outline some of the issues, the problems and
prospects for the use of genetic data in conservation.
THE ENVIRONMENT AND THE DISTRIBUTION OF
GENETIC VARIATION
The earth is not a homogeneous place. This obvious fact
has profound implications for the ways in which organisms
live and evolve and is very probably responsible for much
of the diversity of life around us. Limitations of the extent
of particular habitats and differences in the ways in which
organisms get their livings contribute in part to the large
differences in the amounts and distributions of genetic
variation which we observe. The following sections describe
some of the basics of population genetics theory.
Gene flow and range expansion
One organism’s minor inconvenience to free movement can
be another’s insurmountable barrier. These barriers can be
physical, as for an animal which cannot cross a small
stream, or behavioural, as for a small rodent which refuses
to cross a small gap between patches of forest, or a plant
reliant on a particular species of animal for pollination or
dispersal of its seeds. Behavioural traits can have a large
influence on the distribution of variation within a species.
Even organisms which range over vast areas of ocean can
have very different genetic population structures as a result
of behavioural differences. An example of the extremes
possible are the North American Eel which inhabits streams
along 4000km of coastline and the Humpback Whales of the
North Pacific and North Atlantic Oceans. The eels migrate
to the Sargasso Sea to reproduce as one massive population
and as a consequence the individuals inhabiting streams
show no geographic differentiation. Other fish species
inhabiting the same streams, but which do not leave their
home streams to spawn, show substantial genetic
differentiation. Humpback Whales on the other hand show
genetically distinct subpopulations within ocean basins
despite their ability to roam over huge distances. This
differentiation is apparently the result of female traditions
Part 1. Biological Diversity
in migratory destinations. The eels then show a very high
rate of gene flow while the humpbacks have a low rate of
gene flow among subpopulations, despite ranging over
comparable areas.
These differences in rates of gene flow and population
structuring have major effects on the course of evolution.
A few broad generalisations are possible, though subject to
all sorts of caveats in particular situations. Species
inhabiting large geographic areas and showing high rates of
gene flow show very little or no local differentiation.
Conversely, species with low rates of gene flow are often
divided into distinct populations. At least some of this
distinctness represents adaptation to the local environment.
Adaptations of this sort are familiar to us all in varieties of
crop plants and domesticated animals which, as a result of
artificial selection by humans, perform better in particular
climates and agricultural regimes. Natural selection can
work in a similar way in producing populations with
adaptations to local conditions.
The Earth has only very recently (in geological and
evolutionary terms) emerged from an ice age. This and
other events in the planet’s history have had, and continue
to have, major effects on the nature and distribution of
living things. Much of the northern hemisphere was under
thick ice 10,000 years ago. Most of that ice is now gone
and in its place is forest, prairie, lakes and tundra, all
teeming with life which has managed to colonise these
newly available habitats. Natural processes of change are
still visibly occurring in these regions, suggesting that
populations inhabiting them are not likely to be in genetic
equilibrium. This means that patterns and amounts of
genetic variation reflect historical factors as well as the
present-day situation.
Genetically effective population size
The number of individuals we can count in a population at
any given time can be a surprisingly deceptive measure of
the size of that population in genetic terms. At one extreme
are organisms with vegetative or asexual reproduction such
as aspen where we can stand in a forest surrounded by
genetically identical individuals and a large area can be
populated by only a handful of clones. A number of other
factors commonly found in nature tend to reduce the
genetically effective size of populations below that of the
observed census size. Organisms with limited dispersal
abilities tend to mate with individuals who are more closely
Telated to themselves than the average for the population at
large. This inbreeding reduces the overall genetic variation
of the population relative to what it would have been if
individuals mated at random across the whole population.
Variation in number of offspring produced by different
individuals in a population produces the same effect. If
some individuals have many offspring while others have
few or none the genetic variation of the population is
reduced relative-to what it would have been if everyone had
the same number of offspring. Similarly, populations which
fluctuate in size or pass through a bottleneck of small
population size can also show reduced genetic variation
relative to that expected, all else being equal. Population
geneticists have developed mathematical formulae to take
account of these complicating factors in order to express
population sizes of different organisms in comparable terms
- the genetically effective population size. All these factors,
and others too, can be operating and indicate the
complexities of understanding genetic population structure
of natural populations. We now have the tools with which
to study this structure. Much remains to be done before we
can hope to have a deep understanding of the structures of
natural populations.
One of the most dramatic examples of the potential discord
between our visual impression of a species and its genetic
reality is the Cheetah. This cat was until quite recently
widely distributed throughout Africa and Asia. It has
undergone a severe reduction in its range and numbers but
is still found in widely separated areas of Africa. Recent
surveys of genetic variation in the Cheetah have found
almost no variation - individuals from widely separated
parts of the species range are genetically almost identical.
These results indicate a severe population bottleneck and
subsequent inbreeding. Cheetahs both in the wild and in
captive populations show pronounced effects of inbreeding
not seen in other wide ranging carnivores. This inbreeding
shows itself in reproductive difficulties such as very low
numbers of sperm, many with morphological aberrations,
and high susceptibility to epizootic diseases resulting from
very low amounts of genetic diversity in their immune
systems. The bottleneck responsible for these difficulties
may well have been due to events following the retreat of
the last ice sheet thousands of years ago. Human assaults on
the Cheetah’s range and numbers have certainly not aided
its recovery from the effects of this bottleneck. Similar
effects of inbreeding resulting from recent population
bottlenecks are seen in relictual populations of lions in the
Gir Forest Sanctuary of western India and in the
Ngorongoro Crater in the Serengeti of Kenya.
Outbreeding depression is the converse of inbreeding
depression. If individuals have differentiated genetically
over their range, the mating of individuals from different
parts of that range can result in deleterious effects. This is
presumably because genes from one area do not necessarily
work harmoniously with genes from another area. The
experimental difficulties involved in trying to understand
these effects are great but we do have observations attesting
to their existence in a number of plant species. Several
experiments have demonstrated an ‘optimal outcrossing
distance’, that is fertilization by pollen from distances
greater than the optimum results in reduced fitness, just as
fertilization by pollen from individuals close by can result
in inbreeding depression. There are a few dramatic
examples of outbreeding depression in animal populations.
In Czechoslovakia, Turkish and Nubian Ibex were mixed
with the local Tatra Mountain Ibex and the hybrids were so
poorly adapted that the entire population went extinct.
The genetic effects of habitat alteration and
fragmentation
Human activities cause genetic changes in species by
altering their population structures. Disruption of dispersal
and migration routes and reduction of population sizes are
the most obvious factors. As with natural processes, effects
of particular activities vary depending on the species
considered, some may be affected virtually not at all while
Genetic Diversity
Figure 1.2 Means and ranges of genetic distance between species in selected
vertebrate genera
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0.8 V2 2
Genetic Distance
Oo Mean HMM Range
Note: The numbers of pairwise comparisons of species are in parentheses.
Source: Avise, J.C. and Aguardro, C.F. 1982. A comparative summary of genetic distances in the vertebrates. In: Hecht, M.K., Wallace, B. and
Prance, G.T. (Eds), Evolutionary Biology. Volume 15. Plenum Press, New York.
others may be devastated. For example, many tropical rely on particular species of insects or birds for pollination.
forest trees occur at very low densities over wide areas and Fragmentation of the forest results in very small numbers
Part 1. Biological Diversity
of individuals in each patch. If their pollinators are unable
to cross the gaps between patches, severe inbreeding or
failure to reproduce can result. So even if accidents of
nature do not remove these rare individuals from isolated
patches of forest, they are genetically speaking dead,
despite appearances to the contrary. Other species in the
same isolated patches of forest may maintain large
genetically effective population sizes, either by being
present in higher densities within patches or by having
better dispersal abilities between patches, or both. Of
course, not all species are adversely affected by habitat
fragmentation, especially those dependent on ‘edge’ habitats
such as where forests and open country meet. Species that
thrive in these circumstances range from animals usually
perceived as desirable like White-tailed Deer, to the vectors
of a number of the most devastating human parasites and
diseases.
The effects of small population size depend on the breeding
system of the species and the duration of the bottleneck. If
population size expands rapidly immediately after a
bottleneck, relatively very little genetic variation will be
lost. If the bottleneck lasts for many generations or
recovery is very slow a great deal of variation can be lost.
Of course a population which remains at a very small size
for an extended period is very likely to go extinct as a
result of demographic accidents, probably before deleterious
genetic effects manifest themselves.
If habitat fragmentation eliminates gene flow between parts
of a species’ range these newly isolated populations have
independent evolutionary futures. What this means for the
long-term future for a species is difficult to predict. It is
certainly time to put some serious effort into trying to find
out.
CONCLUSION
The genetic diversity inherent in most species provides the
raw material to respond rapidly to changed circumstances.
This response may not always be adequate and it may not
be in the best interests of humans, as when agricultural
pests and human pathogens develop resistance to our control
measures. Change is, of course, the normal state of affairs
in the living world. What makes our present situation
unique is the rapidity and scale of the change. Our
fragmentation and destruction of habitats constitutes a
massive uncontrolled experiment in ecology and genetics.
We are beginning to understand in outline what needs to be
done to mitigate at least some of the negative effects of this
experiment. Knowledge of the population structures, i.e. the
distribution and amount of genetic variation, of a wide
range of organisms is necessary, as is a much deeper
understanding of the biological significance of different
sorts of variation.
References
Avise, J.C. and Aguardro, C.F. 1982. A comparative summary of
genetic distances in the vertebrates. In: Hecht, M.K., Wallace, B.
and Prance, G.T. (Eds), Evolutionary Biology. Volume 15. Plenum
Press, New York.
Li, W.-H. and Graur, D. 1991. Fundamentals of Molecular Evolution.
Sinauer Associates Inc., Sunderland, Mass.
Contributed by Richard Thomas, Molecular Biology Unit,
The Natural History Museum (London).
2. SYSTEMATICS AND DIVERSITY
This chapter provides a short introduction to systematics:
the branch of biological science responsible for recognising,
comparing, classifying and naming the millions of different
sorts of organisms that exist. As such, systematics provides
the basic framework for the whole of biology, and is the
fundamental discipline of biodiversity. The work can be
divided into a number of activities, including classification,
identification and nomenclature. These are often grouped as
taxonomy, broadly defined as the classification and naming
of organisms. This chapter gives the background for
Chapter 3, which discusses some key theoretical and
practical problems arising from the concept of the species.
BIOLOGICAL CLASSIFICATION
The ultimate task of systematics is to document and
understand the extent and significance of biological
diversity. Within this framework, taxonomy performs four
basic functions: differentiation (recognition of taxa),
identification (universal diagnosis of taxa), symbolisation
(application of universal names), and comparison (relative
Telationships of taxa). Vernacular or folk taxonomies
provide limited local systems for the first three but have
little to tell us about the last.
Individuals and characters are the most basic units of
biological classification. On the basis of features held in
common (attributes or characters), individuals can be
grouped together into a large number of different classes.
These classes are of two kinds (often regarded as sharply
distinct, although in reality they form a continuum). On the
one hand, individual organisms can be divided into such
groups as freshwater, marine, terrestrial, planktonic,
nocturnal, pollinators, etc. Alternatively, they can be placed
into taxonomic categories of species, genera, families,
orders and so on. The former are regarded as artificial
classes, constructed only to serve a particular purpose,
whereas the latter are seen, ideally, as natural groups.
Natural groups comprise individuals with a very large
number of attributes in common, whereas individuals
belonging to artificial groups have relatively few shared
characters. Thus the essential difference between, for
example, ‘marine animals’ and Mammaliais that individuals
of the latter class have far more in common than those of
the former. A natural group, being based on a large number
of characters, can be used for a far wider range of
generalisations and predictions than an artificial group.
Artificial and general classifications are not restricted to
biology. Biology, however, has a unique theory of its own,
the theory of organic evolution. Ideas about evolution can
be divided into a general theory of descent with
modification and special theories about the processes of that
descent (natural selection, neutral theory, etc.). Modern
systematists consider that the general theory of evolution
not only provides a compelling justification for seeking one
natural, general classification for living organisms but also
suggests the basis on which that classification can be most
securely founded: the hierarchic pattern of the ancestor-
descendant sequence, or phylogenetic relationships.
Systematics and Diversity
PHYLOGENETIC RELATIONSHIPS AND THEIR
ESTIMATION
In the past, many biologists have denied that we have
access to sufficient or appropriate information to determine
the phylogenetic relationships of organisms. In the last 25
years, however, spectacular advances in such areas as
molecular biology threaten to overwhelm us with suitable
data. Moreover, during this same period great advances
have also occurred in the theory of systematics and methods
of data analysis.
In an absolute sense, being part of remote history,
phylogenetic relationships cannot be known. What is done
instead is to estimate the most basic feature of the ancestor-
descendant sequence, the pattern of branching points or
nodes. Relationships are defined in terms of common
ancestry. If two species are considered to have a common
ancestor which they do not share with a third species, then
the first two are considered to be more closely related to
each other than either is to the third. This represents the
fundamental three-taxon problem, basic to all phylogenetic
(or cladistic) analysis
Cladistic analysis rests on three basic assumptions: features
shared by organisms (homologies) form a_ hierarchic
pattern; the hierarchic pattern can be expressed by
branching diagrams (cladograms); and the nodes in a
cladogram symbolise the homologies shared by the
organisms subtended by that node (groups). Where data are
in conflict (as they usually are, to a greater or lesser
extent), parsimony is used to find the best supported or
most efficient solution.
Cladistics differs from other methods of classification
because, based on these principles, only special
resemblances are used as evidence of relationship or group
membership. This is in sharp contrast to methods such as
phenetics, in which all resemblances, including character
absences, are regarded as equally informative. Some of the
principles involved here are illustrated in Fig. 2.1.
Cladistics has been at the centre of heated debate, but is
now widely acknowledged to be the best way of
approximating the branching patterns of phylogenetic
history.
FROM HIERARCHY TO CLASSIFICATION
Once a justified hierarchy of phylogenetic relationships has
been established, what relationship should exist between the
hierarchy and classification? Organisms are divided into
kingdoms (animals, plants, etc.), kingdoms into phyla
(Arthropoda, Chordata), phyla into classes (Crustacea,
Mammalia), classes into orders (Decapoda, Rodentia),
families (Cancridae, Muridae), genera (Cancer, Rattus) and
species (Rattus norvegicus, Rattus rattus). Each group
contains the entirety of one or more groups at a lower level.
The categories most often used are shown in Table 2.1, and
see Fig. 2.2. Multiple membership of categories is not
permitted (thus an organism cannot belong to two or more
orders, genera or species at once, with the possible
exception of hybrids).
1. Biological Diversity
Figure 2.1 Establishing the
phylogenetic hierarchy
Characters
Taxa 0 4 2 3 4 5 6 7
MAN - G L i o s v L
ALLIGATOR “ E L s ° ° v t
TUNA FISH - - - - - A v L
SHARK - - - - - T v N
SHARK TUNA FISH MAN ALLIGATOR SHARK TUNA FISH MAN = ALLIGATOR
Notes: Establishing the phylogenetic hierarchy - overall similarity or
special resemblance? In this simple example of a four taxon problem,
the characters are the amino acids (guanine, lucine, serine etc) found
to vary at eight homologous positions in the amino acid sequence of
myoglobin A in man (taxon A), an alligator (B), a tuna fish (C) and a
shark (D) (note that the positions 0-4 in man and alligator have no
equivalent in tune or shark, and that position 0 is also represented in
man). If both presences and absences are counted as equally
informative, the branching diagram on the left most efficiently
summarises the data, but if only presences (special resemblances) are
counted, the cladogram on the right is best. The first solution is that of
the phenetic school of classification, which would continue to accept
the ‘fish’ as a natural group. The second solution is that of the cladistic
school of classification, which would wish to recognise that, in terms
of recency of common ancestry, there is good evidence that the tuna
fish is more closely related to man and alligator than it is to the early-
diverging shark. As a result, the fish is seen to be a paraphyletic
group, of little or no value on natural (phylogenetic) classification (see
also text; based on Patterson, 1980).
Groups which, on the evidence of shared unique characters
(special resemblances), are considered to contain all the
living descendants of a common ancestor are called
monophyletic groups; the mammals are an example. Use of
characters which have evolved more than once leads to the
formation of polyphyletic groups (groups of organisms
which have multiple origins, such as placing birds and
turtles together because they have beaks). If groups are
formed on the basis of unspecialised or non-unique
characters, such as reptiles (which can only be recognised
collectively as members of the amniote vertebrates that are
not mammals or birds), these are termed paraphyletic
groups.
These distinctions are important because they relate to a
continuing debate over the relationship between genealogical
hierarchy (as discovered by cladistic analysis of taxonomic
characters) and formal classification. Most taxonomists
agree that polyphyletic groups once recognised should be
abandoned (although some remain in use, such as
lophophores, a false grouping of the animal phyla
Phoronida, Ectoprocta and Brachiopoda). But many
paraphyletic groups continue to be very widely used, such
as the invertebrates (Metazoa minus Chordata), fish
(Chordata minus Tetrapoda) and Reptilia (Amniota minus
birds and mammals).
Nevertheless, Darwin’s view that our classifications should
correspond to genealogies is becoming more and more
widely accepted. In the last 2-3 decades much progress has
been made in discovering the phylogenetic relationships of
organisms but far more needs to be done. In what follows
it is therefore necessary to appreciate the ideal of hierarchic
classification based on phylogenetic relationships and the
compromise that most existing classifications still represent.
Table 2.1
KINGDOM
DIVISION (Botany) or PHYLUM (Zoology)
CLASS
ORDER
FAMILY
GENUS
SPECIES
Subspecies
The taxonomic hierarchy
Variety (Botany)
Form (Botany)
Note: The categories of the taxonomic hierarchy in descending order
of rank and inclusiveness. There are a few additional less commonly-
used categories, (subphylum, superfamily, tribe, etc.).
Figure 2.2 Basic principles of
classification
ORDER
FAMILY
em Ae Se
p a
f ee
me
4 NS
GENUS GENUS
~\
YA re J.
via \ he
yf Xe a
SPECIES SPECIES SPECIES SPECIES
Notes: Diagram to illustrate the basic principles for turning a scheme
of phylogenetic relationship into a classification (based on Goodrich,
1919, The Evolution of Living Organisms). According to Goodrich,
“individuals are grouped into species, species into genera, these again
into families, orders, classes and phyla, divisions of increasing size and
importance ... the only "fixed points” in a phylogenetic system of
classification are the points of bifurcation, where one branch diverges
from another ... it is here that our divisions should be made.”
Goodrich’s formulation of the problem remains appropriate today,
except that to avoid mis-matches in the ranking of higher categories,
division should proceed from top downward, not from the terminals
(species) upwards. Simple principles of this sort allow us to translate
the phylogenetic hierarchy into a classification hierarchy, although
some compromises and exceptions are still widely accepted in practice,
notably through continued recognition of paraphyletic groups such as
invertebrates, fish and reptiles (see text), and problems created by
hybridization (particularly in plant taxonomy).
TAXONOMIC NOMENCLATURE AND ITS
REGULATION
A separate problem from classification, but often confused
with it, is taxonomic nomenclature. The ultimate goal of
scientific nomenclature is a universal system of
unambiguous names for all recognised taxa. Scientific
names are basic to all biology, and biodiversity is no
exception. In particular, their exact significance has
important implications for conventions, red lists, export
controls, licences or any other legal instruments used to
manage biological diversity.
Taxonomic nomenclature is controlled by voluntary
application of internationally agreed rules or codes.
Separate codes apply to the animal kingdom, plants
(including fungi), and bacteria. In this section the operation
of the zoological code will be outlined and the other codes
briefly compared by noting a few of their differences.
International Code of Zoological Nomenclature
The formation and application of names at the rank of
species (including subspecies), genus and family are
regulated by the Jnternational Code of Zoological
Nomenclature (the Code), and by the use of type specimens.
Cases in dispute are settled through submissions to the
International Commission on Zoological Nomenclature
(ICZN). Names in use below the rank of subspecies (for
polymorphic forms, seasonal variations, hybrids etc.) and
above the rank of superfamily (orders, classes, phyla etc.)
lie outside the scope of the zoological code, and are simply
regulated through usage. This might appear unsatisfactory,
but in practice it gives rise to few difficulties. The major
problems occur with the names of species and, to a lesser
extent, genera and families.
The zoological code depends on two operational principles -
availability and priority - and also governs the formation
of names. To be considered nomenclaturally valid, a species
name must be introduced in combination with a generic
name, and in Latinised form. The species name follows the
generic, never takes a capital, and is usually printed, as
with the generic name, in italics (e.g. Homo sapiens, Rattus
norvegicus, Papilio machaon). If a species is considered
divisible into two or more taxonomically distinct subspecies,
formal trinomens can be introduced. The subspecies
including the population originally described is designated
by tautonymy (Papilio machaon machaon); other subspecies
receive distinguishing third names (Papilio machaon
britannicus).
Availability
For a name of a subspecies, species, genus or family to be
recognised within zoological nomenclature, a number of
requirements must be met. If all these are satisfied, the
name is said to be available; if not, the name is considered
unavailable for the purposes of nomenclature. For a species
name these requirements normally include: a statement that
the name is proposed for a newly recognised species or
subspecies; an indication of how the new taxon differs from
other, related species; and proposal of the name in Latinised
binominal form (i.e. the new species name must be
proposed in combination with a generic name). These are
Systematics and Diversity
some of the basic ingredients of the description, which must
be properly published, in printed form.
Priority
The second basic principle is priority. If what is currently
considered a single species, genus or family has received
two or more available names independently, how would you
choose between them? The basic principle of priority simply
directs that, wherever possible or practical, the oldest or
senior available name must be used. Binominal
nomenclature for animals was first consistently introduced
in the 10th edition of Linnaeus’s Systema Naturae,
published in 1758, and this gives a baseline for priority.
For zoological nomenclature it is therefore unnecessary to
consider names published in any work before 1758 (with
the exception of a single work on spiders published in
1757).
Name, author and date
The two principles of availability and priority come together
in the original published description. It is for this reason
that, when a name is mentioned formally (as in a
catalogue), the original author of the name and year-date of
publication should also be mentioned; thus: Papilio
machaon Linnaeus, 1758.
Types and their function
Species and other taxa are concepts about the organisation
of the natural world, whereas names are artefacts, symbols
intended to designate those concepts. As taxonomic
concepts change, difficulties arise with the application of
existing names. One of the commonest problems occurs
when there are more names available than taxa to be
designated. Which old names apply to which newly
circumscribed taxa? Objectivity in the application of names
is achieved by the use of type specimens.
The code strongly recommends that in original descriptions
the author selects a particular specimen as the type (strictly
holotype) and ensures that it is clearly so labelled and
preserved in a permanent place (normally a museum) so that
it can be studied again in the future. What is the purpose of
such types? It is quite commonly supposed, by those
unfamiliar with biological nomenclature, that the type
specimen represents some sort of ‘standard’ (typical) for
defining the species, perhaps analogous to the standard
metre or standard kilogram used to calibrate rulers or
weights. Nothing could be further from the truth. The type
specimen is simply the name-bearer - it is the specimen to
which the original name is attached. In cases of doubt over
identification with a particular species concept, if you can
decide to which concept the type specimen fits, then the
name automatically follows. Where more than one name is
found to apply, then priority will normally determine which
one is to be used; the other names are synonyms.
Why do names change?
Everyone who makes regular use of biological
classifications soon becomes aware that ‘official’ names can
change. The instability of scientific names is irritating and,
as conservation and wildlife trade legislation becomes more
complex, can lead to real difficulties. Some systematists,
embarrassed that instability gives taxonomy a bad name,
have proposed that a stabilised ‘official list’ should be
1. Biological Diversity
created (for one of the latest rounds of discussion, see
Hawksworth, 1991).
Changes in nomenclature occur for two basic reasons:
problems with names and their application (homonymy,
synonymy, and misidentification, as normally decided by
interpretation of the international code), and revisions of the
system of classification necessary to reflect new scientific
discoveries about taxa and their natural relationships.
Frequently these problems are compounded. While
responsible efforts to avoid ‘unnecessary’ changes brought
about by slavish application of the code are to be
encouraged (because taxonomy is a science to which
nomenclature ought to be subservient), it is futile to imagine
that some fixed, permanently stable list of names can be
drawn up.
To insist on fixity would be far more damaging to
biological science than to accept the minor irritation that, as
our understanding of natural classification changes and
steadily improves, it is necessary to adjust nomenclature
accordingly. However, there are situations where automatic
application of the code can lead to changes considered so
unacceptable that the normal rulings of the code are best set
aside. Such cases are submitted to The International
Commission on Zoological Nomenclature, an international
panel of experts in animal nomenclature whose role is to
decide on the best action in such cases, and then publish
their decisions through the Bulletin of Zoological
Nomenclature.
International Code of Botanical Nomenclature
This code governs the names of fungi as well as green
plants. The ICBN operates in a broadly similar way to the
zoological code, but differs in many details. One obvious
difference is the ‘double citation’ whereby, if there has been
any change in taxonomic assignment or rank of a taxon
since its original proposal, the name is formally to be cited
with the original author’s name in parentheses, followed by
the name of the taxonomist who proposed the change.
Thus the plant known in English as the scentless mayweed
was named by Linnaeus as Matricaria inodora. Later, it
was moved by Schultz-Bipontinus to a separate genus,
Tripleurospermum. This is the accepted name today, and its
authority is formally quoted as Tripleurospermum inodorum
(L.) Sch-Bip. Another difference is that tautonyms are not
permitted for species names. Thus a name like Bison bison,
acceptable under the zoological code, would not be
acceptable in botany. (Tautonymous names below the rank
of species do, however, occur in botany, being created
automatically when plant species are first named; these so-
called antonyms apply to varieties and subspecies.) Unlike
zoological nomenclature, to establish a valid botanical name
it is essential that the original description includes a Latin
diagnosis.
Cultivars are specifically the subject of an additional code,
the International Code of Nomenclature for Cultivated
Plants. Because of biological and other peculiarities, a
number of special provisions also apply to fungi, lichens,
plant hybrids and certain other groupings. One example is
that the name of a lichen is taken to apply to the fungal
part, should it be necessary to consider priority over the
10
application of names to its constituent algal or fungal
elements. At a more fundamental level, there are subtle but
important differences between the botanical and zoological
codes regarding availability and the significance of types.
Changes in the botanical code, and appeals against the strict
application of its provisions, must be directed to the
Nomenclature Section of an International Botanical
Congress, for decision in plenary session.
Codes for the nomenclature of bacteria, actinomycetes,
and viruses
Names for bacteria and actinomycetes are controlled by the
ICNB, the /nternational Code of Nomenclature for Bacteria,
itself controlled by the International Committee for
Systematic Bacteriology. In some respects the bacterial code
is similar to the botanical code (e.g. double citation) but
there are many differences in detail. A_ particularly
important development occurred recently when the
nomenclatural starting date for all bacteria was revised to 1
January 1980, to coincide with publication of the Approved
List of Bacterial Names (Skerman, McGovern and Sneath,
1980).
The names of viruses present exceptional difficulties, and
no international or standard system has been followed.
During the 1966 International Congress for Microbiology
the problem was addressed by an International Committee
on Nomenclature of Viruses (ICNV). This produced a
report, Classification and Nomenclature of Viruses (Wildy,
1971), including recommendations for rules. Since then the
ICNV has become the International Committee on
Taxonomy of Viruses (ICTV), revising and re-revising the
rules and recommendations of Wildy’s report. An almost
complete statement is to be found in Matthews’ (1979)
report, Classification and Nomenclature of Viruses, the
nearest approach yet to an international code for viral
nomenclature.
MAJOR FEATURES OF THE HIERARCHY OF LIFE
The evidence of molecular biology, notably the universality
of the genetic code, strongly favours the idea that all
modern life on Earth is monophyletic.
Ernst Haeckel (1866) was amongst the first to recognise the
enormous diversity of bacteria and other unicellular
organisms, separating many of these life forms (together
with many others that would no longer be included) into a
major group, the Protista, equal in rank to the plants and
animals. This group is no longer formally recognised; some
‘protists’ are currently classified amongst the prokaryotes.
This basal, paraphyletic assemblage comprises the
eubacteria (for which there is good evidence of monophyly)
and archaebacteria (which may or may not form a natural
group). The prokaryotes represent an evolutionary grade in
which DNA is not organised within a nuclear envelope.
The higher organisms, the eukaryotes, form a clade
characterised by possession of a double nuclear membrane.
The eukaryote clade includes the three major groups of
macro-organisms, the green plants, fungi and animals,
together with many unicellular and other simple organisms
now often referred to as ‘protists’. The protists include the
myxomycetes (slime moulds), protozoans and various
groups of algae, including green algae (chlorophytes),
chromists (chrysophytes, golden brown algae etc.), and
rhodophytes or red algae; the chlorophytes form a
monophyletic group with the green plants (Bremer, 1985).
Plants
A major group, comprising the green algae and the land
plant kingdom, can be recognised as a natural group. Of
three primary divisions, the Chlorophyta (green algae)
comprise a complex paraphyletic group from within which
the land plants (embryophytes) have arisen. The most basal
groups of land plants are the liverworts and hornworts, and
then the mosses. The next level of organisation is
represented by the tracheopytes (characterised by the
possession of vascular tissue), including lycopods, horsetails
and ferns. Beyond this level are the seed plants
(spermatophytes), including cycads, Ginkgo, conifers, a
group comprised of Ephedra, Gnetum and Welwitschia, and
finally the flowering plants (angiosperms). The angiosperms
are a vast and complex assemblage, traditionally divided
into the monocotyledons (probably monophyletic) and the
dicotyledons (paraphyletic).
Fungi
The fungi form a major kingdom, divisible into the
Oomycetes and the true fungi, the Eumycota. According to
Tehler (1988), the true fungi (identifiable as a natural group
on the basis of 25S RNA and chitin cell walls) can be
divided into four divisions, one of which includes the
Dicaryomycotina. The dicaryomycetes are themselves
divided into three classes: the Ascomycotina (moulds,
yeasts), Protobasidiomycotina, and Basidiomycotina(smuts,
trusts, bracket fungi, mushrooms, toadstools). A number of
poorly-known fungal groups probably do not fit into this
scheme, but the ‘fungi-imperfecti’ (Deuteromycotina) are an
unnatural assemblage of forms (including many moulds)
unknown in their sexual stage, most of which are believed
to be non-sexual stages of ascomycetes and basidiomycetes.
Animals
The higher, multicellular animals (Mesozoa and Metazoa)
are usually regarded as monophyletic, the principal basal
members being the mesozoans and poriferans (sponges),
followed by coelenterates (jelly fish and cnidarians) and
platyhelminths (flatworms). The molluscs, arthropods
(including insects), echinoderms (starfish, sea urchins etc.)
and vertebrates are conventionally grouped together at the
apex of the animal hierarchy.
In conclusion, although some major features are discernible,
our knowledge of the hierarchical pattern of life, even at
this most general level, appears very limited. However,
new molecular evidence, such as the 18S rRNA data studied
by Lake (1990) and others, holds the promise of yielding
far greater understanding. Margulis and Schwartz (1988)
should be consulted for further information on all the
recognised phyla of organisms, their biology, relationships
and taxonomy.
SYSTEMATICS AND THE MEASUREMENT OF
BIODIVERSITY
Ecologists have measured diversity either by estimating
11
Systematics and Diversity
species richness (number of species) in an area, or by one
or more indexes combining species richness and relative
abundance within an area. Some attempts have also been
made to measure change in species richness (species
turnover) between areas. These solutions to the problem of
measuring biodiversity are limited because species richness
takes no account of the differences between species in
relation to their place in the natural hierarchy, and because
relative abundance is not a fixed property of species,
varying widely from time to time and place to place.
Furthermore, in many environments most taxa are virtually
or even completely unknown.
For some time conservationists have called for a
measurement of diversity more clearly related to overall
genetic difference. For example, regarding the problem of
differential extinction, IUCN/UNEP/WWF (1980) noted
that "the size of the potential genetic loss is related to the
taxonomic hierarchy because ... different positions in this
hierarchy reflect greater or lesser degrees of genetic
difference ... the current taxonomic hierarchy provides the
only convenient rule of thumb for determining the relative
size of a potential loss of genetic material."
Measurements of diversity are now being proposed that
either attempt to measure genetic difference directly, or
indirectly through use of the taxonomic (cladistic) hierarchy
(Williams er al., 1991; Faith, in press). Apart from
scientific debate still not fully resolved, the latter approach
is more practical because we already have a "rule of
thumb" taxonomic hierarchy (which is being steadily
improved through the application of cladistic analysis,
notably to molecular data), whereas reliable estimates of
overall genetic differences between taxa are virtually non-
existent.
Based on the shared and unshared nodes between taxa
(equivalent to position in the taxonomic hierarchy), a
number of taxonomic diversity indices have now been
developed. Of these, the most distinct are root weight,
higher taxon richness and taxonomic dispersion. The first
places highest individual value on taxa which separate
closest to the root of the cladogram and comprise only one
or relatively few species; in effect this gives high weighting
to relict groups. Higher taxon richness favours taxa
according to their rank and number of included species.
Dispersion, the most complex of the measures proposed so
far (Williams er al., 1991), endeavours to select an even
spread of taxa across the hierarchy, sampling a mixture of
high, low and intermediate ranking groups. See Fig. 2.3 for
illustration of these concepts.
For a given group these measures, together with simple
species richness if desired, can be used to compare the
biotic diversity of any number of sites. The measures can
also be expressed as percentages. Thus a site with viable
populations of all species in a group would have a diversity
score of 100%, while a site without any species of the
group in question would score zero. In reality, of course,
most sites have only a selection of species, and so receive
various intermediate scores.
Such assessments allow us to compare all sites with each
other, and rank them individually from highest to lowest
1. Biological Diversity
Figure 2.3 Measures of biodiversity
SHARK
SALMON
TURTLE
BIRD
SNAKE
Notes: The practical need for measures of biodiversity. Assume there
is a small zoo keeping six species of vertebrates: a shark, a bony fish
(salmon), a rat, a turtle, a bird and a snake but only half can be
maintained in future (each one costs the same). If the objective is to
display as ‘good’ a sample of biodiversity as possible, which three
should be selected? Accepting the phylogenetic relationships in the
diagram, species richness offers no help - all 20 possible choices are
the same. Taxic diversity measures will help us choose, but the result
will be dependent on which index we use. Root-weight selects shark,
bony fish and rat. Higher taxon richness selects a shark, bony fish plus
one of the remainder. Taxic dispersion chooses shark and rat plus bird
or snake. Dispersion is probably the criterion that corresponds most
closely to an intuitive notion of diversity.
diversity. However, if we then take some action on this
(such as conserving a particular site), the same measures
are unlikely to be directly comparable for making a second
decision (such as choosing a second conservation site). This
is because, in most real situations at least, there will be
considerable overlap in the presence of species at particular
sites.
In a seminal work on the measurement of diversity,
Whittaker (1972) introduced the concepts of alpha, beta and
gamma diversity. The measurements just described, giving
diversity values for single sites, are examples of alpha
diversity. The beta and gamma diversity concepts relate to
changes in diversity between sites at local (beta) and
geographical (gamma) scales. An essential part of these
relational concepts is the idea of species turnover - the
degree to which species present at one site are replaced by
others at different sites. For use in assessing the relative
value of multiple sites for the conservation of biodiversity,
the idea of species turnover is translated into the principle
of complementarity, implemented in combination with a
taxonomic diversity index. This is returned to in
Chapter 15.
References
Bremer, K. 1985. Summary of green plant phylogeny and
classification. Cladistics 1:369-385.
Faith, D. (in press). Conservation evaluation and phylogenetic
diversity. Biological conservation.
Goodrich, E.S. 1919. The Evolution of Living Organisms. Jack and
Nelson, London.
Haeckel, E. 1866. Generelle Morphologie der Organismen, 2. Berlin.
Hawksworth, D.L. (Ed.) 1991. Improving the stability of names: needs
and options. Koeltz Scientific Books, Koenigstein. (Regnum
Vegetabile 123).
International Code of Nomenclature for Bacteria. 1975. American
Society for Microbiology, Washington.
International Code of Botanical Nomenclature. 1988. International
Association for Plant Taxonomy (Europe).
International Code of Nomenclature for Cultivated Plants. 1980.
International Commission for the Nomenclature of Cultivated
Plants, IUBS.
International Code of Zoological Nomenclature. 1985. International
Trust for Zoological Nomenclature, London.
TUCN/UNEP/WWF 1980. World Conservation Strategy. living
resource conservation for sustainable development. Gland,
Switzerland. .
Lake, J.A. 1990. Origin of the Metazoa. Proceedings of the National
Academy of Science USA 87:763-766.
Margulis, L. and Schwartz, K.V. 1988. Five Kingdoms: an illustrated
guide to the phyla of life on earth. W.H. Freeman, New York.
Matthews, R.E.F. 1979. Classification and nomenclature of viruses.
Third report of the International Committee on Taxonomy and
Viruses. Intervirology 12:131-296.
Patterson, C. 1980. Cladistics. Biologist 27:234-240.
Skerman, V.D.B., McGowern, V. and Sneath, P.H.A. (Eds) 1980.
Approved list of bacterial names. International Journal of
Systematic Bacteriology 30:225-420.
Tehler, A. 1988. A cladistic outline of the Eumycota. Cladistics 4:227-
277.
Whittaker, R.H. 1972. Evolution and measurement of species diversity
Taxon 21:213-251
Wildy, P. 1971. Classification and nomenclature of viruses. In:
Melnick, J.C. (Ed.), Monographsin Virology 5. London: Academic
Press, London.
Williams, P.H., Humphries, C.J. and Vane-Wright, R.I. 1991.
Measuring biodiversity: taxonomic relatedness for conservation
priorities. Australian Systematic Botany, 4:665-679.
Williams, P.H. (unpublished). Afrotropical antelopes - priority areas
for biodiversity. Progress report to the Natural History Museum,
London, WCMC and IUCN-SSC.
Abridged from a document written by R.1. Vane-Wright,
Biodiversity Programme, The Natural History Museum
(London).
3. SPECIES CONCEPTS
An understanding of the species concept is basic to an
understanding of biological diversity because species are
almost universally used as the units in which diversity is
measured.
WHAT IS A SPECIES?
This simple question has troubled biologists for more than
two centuries. Although accepted so widely as a ‘natural’,
basic or fundamental unit, many conflicting definitions of
species have been coined, and agreement is still lacking.
The range of definitions reflects, to a large degree, the
differing interests and differing theories of individual
scientists about the origin of diversity itself - literally from
Genesis to Darwin and DNA. This process has not stopped,
continuing for example with the debate over the importance
of neutralism or selectionism in the evolutionary process.
Furthermore, many scientists have entered the debate from
practical knowledge of particular groups of animals or
plants. As there are major differences in the biology of
different groups, with consequent variations in the patterns
and processes of species formation, it is hardly surprising
that species and species concepts are heterogeneous both in
theory and practice.
One of the most fundamental aspects of the problem is
variation. Most if not all animals and plants show variation,
every individual often being demonstrably unique. Within
a population variation can be continuous (such as height or
weight) or discontinuous (such as sex or handedness),
environmental in origin (such as human language) or
genetic (such as blood group). Variation can also be seen in
time between successive generations (seasonal variation),
and in space across allopatric populations (geographical
variation: clines, demes, races and subspecies).
The species problem is, in part, a history of how biologists
have tried to manage this problem of variation. In
particular, how can we classify variable organisms into
discrete groups, tempered by knowledge of the existence,
origin and maintenance of that variation? Modern species
concepts divide into two main groups, those concerned with
process and those concerned with pattern. We thus need to
examine the processes of segregation, isolation and
recognition responsible for the differentiation and cohesion
of populations, and the patterns we perceive through
comparison of the products of those differentiation
processes.
EARLY SPECIES CONCEPTS
The word ‘species’ literally means outward or visible form.
Conspicuous natural species have long been recognised by
people of many local cultures. With the emergence of
natural science in the 17th and 18th centuries, attempts
were made to catalogue the whole of biological diversity, in
all its manifestations and variations. Early approaches to
dealing with the species problem were influenced by two
very different philosophical views, essentialism and
nominalism. In practice, however, both were usually
abandoned in the face of increasing empirical knowledge of
the life cycles of organisms and how they reproduce.
Species Concepts
According to the typological species concept, based on
essentialist principles, which was widely adopted during
much of the 18th and 19th centuries, every organism
corresponds to some idealised plan. The task of the
taxonomist involved recognising each fundamental design,
and describing, diagnosing or divining the essential features
of those designs or ‘types’, so that individual organisms
could be assigned to them.
In practice this often led to arbitrary divisions. Very
different plants or animals were often lumped together
because they shared certain ‘essential’ features; this was
particularly evident amongst higher taxa, such as Linnaeus’s
group Vermes. By the same token, what we would now
recognise as different forms of one and the same animal or
plant were often separated because they conformed to
different idealised types - in its most extreme manifestation,
in many higher taxa two different sexes exist which
according to this view could be classified as separate
species, plainly a nonsensical view.
The most extreme opposing view states that only individuals
exist in nature. Taxonomic groups are seen as man-made
abstractions allowing us the convenience of being able to
refer to large numbers of individuals collectively, and
nothing more. They have no objective or independent basis
but are merely convenient ‘pigeon-holes’ for dividing up or
handling diversity.
Few scientists now accept that this nominalist approach is
applicable to species, but it is still widely considered to
apply to higher taxa. Most cladists and other taxonomists
concerned with natural classification deny nominalism at all
levels - the kingdom is seen as ‘real’ as the species
(Loevtrup, 1987). Some cladists, however, deny reality to
the species level, seeing species as only something in the
process of becoming, while higher taxa are considered
permanent real entities. With such deep divisions in the
philosophical views of taxonomists, it is hardly surprising
that there is still no agreement over the species concept.
EVOLUTIONARY THEORY AND POLYTYPIC
SPECIES
Evolution and genetics
Following the emergence of Darwinism in the 1860s, and
the general acceptance of the theory of evolution, the
typological approach began to be questioned. Darwin
himself suggested that “our classifications will become, so
far as possible, genealogies". To Darwin, species were no
different from other taxa (a view currently advocated by
Nelson, 1989), and he expressed relief at being freed from
"the vain search for the undiscovered and undiscoverable
essence of the term species". Darwin, however, had no
reliable theory of inheritance. With the development of
genetics and population biology, including statistics,
scientists began to develop rational explanations for the
origin and inheritance of variation, and apply this
understanding to a radically different view of the nature of
taxa - and species in particular.
1. Biological Diversity
Polytypic species
One of the first major impacts of population thinking on
taxonomy was the concept of polytypic species. According
to this idea, many widespread species-level taxa show more
or less discontinuous geographical variation describable by
the use of trinomens, or subspecies. Previously such
variation was recognised haphazardly by the occasional
naming of ‘varieties’ or, alternatively, by the description of
increasingly large numbers of allopatric species, many of
which differed only in details of coloration or other
superficial characters. Such patterns were seen to reflect
both common ancestry and local adaptation, as species were
thought to spread from their geographical places of origin
and differentiate under the influence of natural selection.
Subspecies were seen virtually as ‘species in the making’.
This approach, including the trinominal nomenclature
(genus, species and subspecies), was preadapted to become
the basis of an influential new vision of the species.
THE BIOLOGICAL SPECIES CONCEPT
The biological species concept is particularly associated
with the work of three zoologists, Theodozius Dobzhansky,
Julian Huxiey and Ernst Mayr. This view concentrates not
on logical classes or plans but on the idea of the species as
a process, a closed reproductive community or breeding
system. According to Mayr (1969), species are groups of
interbreeding [or potentially interbreeding] natural
populations that are reproductively isolated from other such
groups. The basic idea of a biological species is that of a
‘pool’ of genes available for re-combination through sexual
reproduction, but not with genes belonging to other gene
pools, from which they are ‘protected’ by a variety of
Tecognition and isolation mechanisms (behavioural,
physiological, genetical, etc.). Thus the biological species
to which a given individual belongs is determined by the
limits of the populations with which it interbreeds, or
potentially interbreeds.
The biological species concept, or some variant of it, is
probably the most widely accepted view of the species held
by biologists today. Extreme versions of the concept, such
as Huxley’s (1940) definition of species as "distinct self-
perpetuating units with an objective existence in nature, and
therefore on a different theoretical footing from genera or
families or other higher categories" approach the
evolutionary species concept, in which species are seen as
the fundamental units of evolution (rather than haphazard
by-products of it).
Recently, certain proponents of the biological species
concept have split into two ‘camps’: those supportive of the
idea that species distinctness is mainly brought about and
maintained by selection for isolating mechanisms (isolation
concept), and those emphasising greater importance for
inherent mate-recognition systems in this role (the
recognition concept). The debate has led to further
proposals, such the cohesion concept. According to
Templeton (1989) this idea draws on all three major
variants of the biological species (the evolutionary, isolation
and recognition concepts), and defines species as the "most
inclusive population of individuals having the potential for
phenotypic cohesion through intrinsic cohesion
14
mechanisms". The intrinsic mechanisms relate to gene flow
and ecological equivalence.
All variations of the biological species concept suffer from
a number of practical shortcomings and limitations. They
are inapplicable to the very large number of animals and
plants that reproduce with only irregular genetic
recombination, or without it altogether (asexual or agamo-
species). In sexually reproducing species the limits of
genetic re-combination are rarely known and have to be
inferred from indirect evidence, and there is further
uncertainty regarding species limits when the concept is
applied over wide geographical ranges or over time.
Superspecies and syngameons
As already noted, the biological species concept was
developed by zoologists from the idea of grouping allopatric
(not overlapping geographically), modestly differentiated
races or subspecies into single polytypic species. This
system was elaborated to include a further concept, that of
the superspecies, consisting of assemblages of more
strongly differentiated groups of populations, or
semispecies. Semispecies have geographically
non-overlapping but contiguous (parapatric) distributions,
permitting gene exchange at their boundaries. Most
Significantly, they are seen as ecological equivalents, and
thus unable to coexist as stable, fully differentiated species.
Following Turesson, botanists have long recognised a
related concept, the syngameon, whereby groups of
sympatric (geographically overlapping) semispecies coexist.
Gene flow may be slight or extensive, and their continued
existence depends on ecological vicariance, occupying stable
and distinct local habitats (such as contiguous forest and
open formations). If such a patchy environment is destroyed
and replaced by a different ecosystem, the separate
semispecies usually fuse through hybridisation.
The advent of genetic fingerprinting techniques has now
permitted zoologists to appreciate that gene flow between
more or less closely related but perfectly ‘good’ sympatric
species of animals may be commonplace. One of the most
recent discoveries of this kind is reported by Templeton
(1991), who quotes work showing that significant gene
exchange can occur between Bison and certain species of
Bos (domestic cattle). This example demonstrates that
species sufficiently distinct to have been placed in different
genera can have this type of relationship, empirically
violating the most basic tenet of the biological species
concept, the separateness of gene pools.
Tokogenetic and phylogenetic relationships
In order to understand continuing disagreements over the
significance and definition of species, it is necessary to
appreciate that two quite separate goals are being pursued.
Species serve as the basis for describing and cataloguing the
elements of biodiversity, and in our attempts to discover the
historical relationships of those diverse elements. Species
are also widely regarded as fundamental units of evolution,
being both the products of speciation and the things which
are thought to speciate. Thus the single word, species,
serves the needs of systematics (discovery of empirical
patterns) and the needs of population biology (formulation
of process theories). Once the existence of these two
separate goals is acknowledged, it becomes easier to make
sense of the multiplicity of species concepts, many of which
represent only differences of emphasis within the two major
divisions.
Another way to think about this problem is to consider two
major sorts of genetic relationships: those between
individuals (tokogenetic, or blood relationships) and those
between taxa (phylogenetic, or historical relationships).
What is truly unique about species may simply be that they
lie at the junction of both types of relationship (Nixon and
Wheeler, 1990). Higher taxa, and their interrelations,
represent a fixed, historical past. Below the species, at the
level of demes and populations, all is change, with mutation
and genetic recombination affecting every new life cycle,
every generation of individuals. Species exist at a dynamic
limit between the two, with tokogenetic processes
maintaining cohesion yet allowing change, while historical
accidents fragment species into separate phylogenetic
lineages. Such ideas form the basis of yet another species
concept, that of phylogenetic species.
THE PHYLOGENETIC SPECIES CONCEPT
According to this view, species are irreducible clusters of
organisms diagnosably distinct from other such clusters, and
within which there are parental networks of ancestry and
descent. Nixon and Wheeler (1990) have defined the
concept as “the smallest aggregation of populations (sexual
reproduction) or lineages (asexual reproduction) diagnosable
by a unique combination of character states in comparable
individuals".
This view of species places the emphasis not on
reproductive process but on the most general aspect of
taxonomic diversification, that of differentiation. In some
cases differentiation results in reproductive isolation but in
many cases it does not. Thus the existence of reproductive
isolation is evidence of diagnostic characters but new
characters which become fixed within a population do not
necessarily affect reproductive isolation.
An inherent danger in such a view is that, by reductio ad
absurdum, every population, stage, morph or even
individual organism could be elevated to separate species
status. For this type of definition to be operational it would
also be essential to emphasise the critical importance of
reproductive community, or cohesion, more or less in
Templeton’s sense. Even then, a consequence of applying
the phylogenetic species concept, compared with the
biological species concept, would be a very large increase
in the number of species recognised (Nelson and Platnick,
1981).
SPECIES IN PRACTICE
Empirical consequences of different concepts
Cracraft (1989) has provided some examples of the striking
differences that can arise in evolutionary and taxonomic
conclusions, dependent on the species concept applied.
Cracraft’s examples all concern parapatric birds of
15
Species Concepts
debatable specific or subspecific status, with evidence of
hybridisation in contact zones. His cladistic analyses suggest
that many biologically defined ‘subspecies’ that hybridise on
contact are less closely related to each other by descent than
they are to other, full ‘species’ with disjunct distributions.
Thus, as accepted under the phylogenetic species concept,
species separable on phylogenetic criteria may be
interfertile, while polytypic species recognised on biological
(interbreeding) criteria may not be the ‘units of evolution’.
At the practical level, these alternative approaches give rise
to major differences in the classification and status given to
populations and groups of populations. As already noted,
the phylogenetic concept or approach leads to the
recognition of far more species (and fewer subspecies) than
the biological species concept. In terms of formal
classification, it lacks the major practical advantage of
trinomens - we would tend to lose sight of the wood for the
trees.
Subspecies
Many species of geographically variable and conspicuous
organisms, such as birds, have been subdivided into
numerous subspecies. Butterflies, for example, are thought
to comprise about 17,500 full species, but the number of
currently recognised subspecies approaches 100,000. Many
of these subspecific taxa (particularly those from small
islands or isolated mountains) are fully diagnosable - that is,
virtually every individual can be reliably identified to
subspecies, regardless of knowledge of where it was found.
Such subspecies would qualify as species under a
phylogenetic species concept.
On the other hand, this is not true for all so-called
subspecies, notably many of those described from large
islands or continental areas. In many of these cases
subspecies are only recognised on a statistical basis, so that
individuals cannot be reliably diagnosed, and only identified
with the aid of knowing where they came from. Typically,
this represents the phenomenon of clinal geographic
variation. At the extreme, the most distant populations in
long clines may be so distinct that in areas of overlap they
may behave as separate biological species and be fully
diagnosable locally (rassenkreis and ring species: Mayr,
1963). Even in less extreme situations, the opposite ends of
a cline may be more strikingly distinct than related, fully
diagnosable subspecies, or even full species.
The implications of this are that for the assessment of
biodiversity there is no easy answer to ‘the subspecies
problem’ any more than there is to the species problem.
Species status bears no direct or simple relationship to
degree of phenetic differentiation, or to any measure such
as genetic distance. Species (and subspecies) are determined
by relational properties, not by absolute criteria, be they
essences, reproductive mechanisms or distance measures.
The state of the science
At the broadest scale, we know very few organisms well
enough to consider the subtle, albeit highly significant,
interpretations that such insights as the phylogenetic species
concept or the syngameon might lead us to consider. In
1. Biological Diversity
particular, the vast majority of named species are known
only from morphology and limited knowledge of their
geographical distributions. For these species we know
Virtually nothing about their individual breeding systems,
gene flow, ecology or even, in most cases, their cladistic
relationships. Such species are often referred to as
morphospecies.
The present state of taxonomy, carried out by different
scientists working at different times to different theories and
philosophies and on imperfectly known groups of widely
differing size, taxonomic apparency and life-cycle
characteristics, ensures that species currently recognised are
not comparable entities. Following the successive rise of
population biology and phylogenetic systematics, there is
some prospect if not of harmonising species concepts at
least of clarifying what is meant by a particular scientist in
a particular context.
CONCLUSION
For the present we have to manage with a very imperfect
and inconsistent system of classification, even at the
supposedly fundamental level of species. In practice we
have not advanced much beyond the position outlined long
ago, that a species is what a competent systematist says it
is (Regan, 1926). Although much can and should be done
to improve this state of affairs, a lack of certainty should be
accepted as inherent to the subject.
However, this strong limitation on the use of species as
comparable units is all too often forgotten when species
numbers are handled in aggregate, as with many practical
conservation issues or theoretical discussions of
biodiversity. Conclusions reached on this basis run a risk of
being inaccurate, spurious or even completely misleading.
16
If species, instead of being treated like independent and
equivalent units of diversity, are placed in their proper
relational context of the entire hierarchical classification,
some of the problems caused by this limitation can be
avoided.
References
Cracraft, J. 1989. Speciation and its ontology: the empirical
consequences of alternative species concepts for understanding
patterns and processes of differentiation. In: Otte, D. and Endler,
J.A. (Eds), Speciation and its Consequences. Sinauer, Sunderland,
Mass. Pp.28-59.
Huxley, J.S. 1940. Introductory: towards the new systematics. In:
Huxley, J. (Ed.), The New Systematics. Oxford University Press,
London. Pp.1-46.
Loevtrup, S. 1987. On species and other taxa. Cladistics 3:157-177.
Mayr, E. 1963. Animal Species and Evolution. Harvard University
Press, Cambridge, Mass.
Mayr, E. 1969. Principles of Systematic Zoology. McGraw-Hill, New
York.
Nelson, G. 1989. Species and taxa: systematics and evolution. In:
Otte, D. and Endler, J.A. (Eds), Speciation and its Consequences.
Sinauer, Sunderland, Mass. Pp.60-81.
Nelson, G. and Platnick, N. 1981. Systematics and biogeography:
cladistics and vicariance. Columbia University Press, New York.
Nixon, K.C. and Wheeler, Q.D. 1990. An amplification of the
phylogenetic species concept. Cladistics 6:211-223.
Regan, C.T. 1926. Organic evolution. Report of the British Association
for the Advancement of Science 1925:75-86.
Templeton, A.R. 1989. The meaning of species and speciation: a
genetic perspective. In: Otte, D. and Endler, J.A. (Eds), Speciation
and its Consequences. Sinauer, Sunderland, Mass. Pp.3-27.
Templeton, A.R. 1991. Genetics and conservation biology. In: Seitz,
A. and Loeschcke, V. (Eds), Species Conservation: a population-
biological approach. Birkhauser, Basel. Pp.15-29.
Text written by R.I. Vane-Wright, Biodiversity Programme,
The Natural History Museum (London).
4. SPECIES INVENTORY
The objective of this section is to explore how far global
biodiversity may have been accounted for by taxonomic
description, emphasising diversity at the species level. This
is done with reference to the total number of species
currently recognised (itself very imprecisely known) and the
degree to which we can estimate the completeness of
taxonomic knowledge.
Existing knowledge of geographical and other variation in
species richness provides a useful starting point, but this
knowledge is heavily biased. Unfortunately, our
understanding of the best-known taxonomic groups and
best-known parts of the world remains an insufficient basis
for predicting more general patterns, or for rigorously
testing explanations for such patterns as have been
identified. Any estimation that may be made of the overall
extent of global species richness remains staggeringly
imprecise. Even so, for at least multi-celled animals and
green plants, and perhaps for all eukaryotes (i.e. all of life
except for microorganisms such as bacteria) it is possible to
predicate useful lower and (with less confidence) upper
limits to the extent of regional and global species richness
of the major groups. Also, it is now reasonably clear just
what the major gaps in our understanding are, so that we
have a good idea of which new data are needed to improve
on present estimates. Work in progress that involves
intensive sampling of species-rich groups (e.g. insects) in
especially species-rich areas (e.g. moist tropical forests)
promises to provide a much more reliable picture of major
global species richness patterns and a more reliable basis
for estimating the number of species with which we share
the planet.
It must be emphasised that data discussed in this section that
may be pertinent to species richness estimates should not,
in the current poor state of knowledge, be applied directly
to estimations of possible species extinction rates via loss or
degradation of habitat. Existing data on range sizes,
patchiness of distribution and population structure of the
poorly-known organisms discussed here are such that no
direct connection between numbers of species present at one
site or in one region and the threat posed to the continued
existence of any one of those species by the loss of a given
area of habitat can be made.
CURRENT STATUS
Here we consider how many extant species of organisms
have already been described and assess at what rate the
existing inventory is growing and improving.
The number of described species
The number of species which have been described and the
number currently regarded as valid are not precisely known
for many groups of organisms. For the best known groups,
all of which are relatively small (e.g. birds with 9,881
species, Sibley and Monroe, 1990), catalogues and counts
are very complete. Variations in published figures are
largely because of differences in whether certain taxa are
regarded as ‘good’ species or not. Accurate figures for
currently recognised species are also available for some
17
Species Inventory
groups (e.g. bacteria with 3,058 recognised species as of
1991) in which it can be assumed a major proportion
remains undescribed. Much improved counts have recently
become available for some substantially larger groups, such
as the vascular plants (260,000 species in total) and fungi
(70,000 species). Counts for animal groups with many
described species, as Tables 4.1 and 4.2 illustrate, mostly
remain much less precise. Disparities between the various
figures very recently furnished for individual groups such
as the molluscs, annelids and platyhelminths (Table 4.1) and
Diptera (Table 4.2) are particularly striking. On
investigation, only some of the apparent discrepancies turn
out to be because of differences in the year up to which
counts had been made; others appear to result from
confusion between the number of nominal species (i.e. all
species that have ever received a separate name no matter
what their current status) and the often much lower number
of species recognised as valid, as well as from simple
miscalculation or oversight. Some of the largest groups
(e.g. the insect orders Coleoptera and Diptera) are, in fact,
relatively well catalogued, but animal taxonomists have
tended to place little emphasis on providing accurate tallies
of described species that are regarded as valid at any
particular point in time. Largely as a result, figures for the
biota as a whole that have been published in recent years
vary considerably, from around 1.4 million to more than
1.8 million. This imprecision is far exceeded by that
involved in the attempts to estimate total species richness
(including as yet undiscovered and undescribed species)
discussed below, but it is in some respects more surprising.
Estimates for numbers of currently recognised and
described species are given here, mostly rounded to the
nearest five thousand, for the groups that make the largest
contributions (see Table 4.3 and Fig. 4.5) - but without any
pretence to high accuracy. In arriving at these figures
relevant specialist opinion, as well as the most recent
literature, was taken into account. Including all of the
smaller groups not listed in Table 4.3, the overall figure
reached is approximately 1.7 million. A more accurate
count is likely to produce a somewhat higher figure.
Deficiencies of the existing database
The evidently low priority accorded by taxonomists to
keeping track of how many species have been described
stems in large measure from the knowledge that the
biological significance of these data is slight. For all but the
best known groups, if a species count is a measure of
anything it is of taxonomic effort expended, and this is
clearly seen to be arbitrary by most biological criteria. Even
in terms of the taxa that ostensibly have been dealt with by
the descriptive process much uncertainty exists, as
catalogues of described species, however carefully
compiled, include the results of poor taxonomy as well as
good. When careful reassessments (taxonomic revisions) are
made, it is common to find that a relatively high proportion
of previously recognised ‘species’ are not, in fact, distinct.
To put it in taxonomists’ jargon, most parts of the existing
inventory contain substantial amounts of unrecognised or at
least unreported synonymy. In some groups, further
imprecision arises from a fundamental lack of agreement as
to just what constitutes a species.
1. Biological Diversity
Table 4.1 Estimated numbers of described extant species in major animal groups
Mayr et a/. Barnes May May Brusca & Brusca
(1953) (1989) (1988) (1990) (1990)
‘Protozoa’ - - 260,000 32,000 35,000
Porifera 4,500 5,000 10,000 - 9,000
Cnidaria 9,000 9,000 10,000 9,600 9,000
Platyhelminthes 6,000 12,700 - = 20,000
Rotifera 1,500 1,500 - - 1,800
Nematoda 10,000 12,000 1,000,000 ? - 12,000
Ectoprocta 3,300 4,000 4,000 - 4,500
Echinodermata 4,000 6,000 6,000 6,000 6,000
Urochordata 1,600 1,250 - 1,600 3,000
Vertebrata 37,790 49,933 43,300 42,900 47,000
Chelicerata 35,000 68,000 63,000 - 65,000
Crustacea 25,000 42,000 39,000 - 32,000
‘Myriapods’ 13,000 10,500 - - 13,120
Hexapods 850,000 751,012 1,000,000 ? 790,000 827,175 +
Mollusca 80,000 50,000 100,000 45,000 100,000 +
Annelida 7,000 8,700 15,000 - 15,000
Notes: ‘Protozoa’, a paraphyletic group, is used in the ‘traditional’ zoological sense. The ‘Myriapods’ consist of the Chilopoda (centipedes) and
Diplopoda (millipedes) together. Apart from two exceptionally high figures - those for Protozoa and Nematoda - provided by May (1988), who
presumably intended these as estimates of actual rather than described species, most estimates, even the highest of a range for any given group,
are probably conservative. The extent of the great variation in totals for some groups is inexplicable. For example, while Brusca and Brusca (1990)
suggest 100,000+ as a likely figure for described species of molluscs a totalling of the figures given for the individual mollusc classes by the same
authors provides a total of around 50,000.
Table 4.2 Number of described species in the four major insect orders
Southwood Arnett May Brusca & Brusca
(1978) (1985) (1988) (1990)
Coleoptera 350,000 290,000 300,000 300,000+
Diptera 120,000 98,500 85,000 150,000
Hymenoptera 100,000 103,000 110,000 125,000
Lepidoptera 120,000 112,000 110,000 120,000
Notes: Some recent estimates of the number of described species in the four major insect orders. An accurate figure for Hymenoptera is probably
not very different from any of the fairly consistent estimates shown, while one for Diptera probably lies towards the middle of the very wide range
indicated here. The estimates for Coleoptera and Lepidoptera, on the other hand, are probably all far too low.
The existing inventory of described species may, as at around 300,000 species (Table 4.4), although thorough
discussed below, provide a poor basis for estimating the counts of a sample of coleopterous families suggest that
true extent of global species richness. However, where 400,000 is a likely minimum. If the figure of 300,000 does,
extrapolative methods are adopted that do involve the use of in fact, represent an underestimate of the order of 100,000
described species counts, the accuracy with which the species its use in the calculations outlined above would lead
counts have been made will often have a substantial to underestimation of the biota as a whole by as much as
influence on results. For example, let us assume that our five million.
approach to estimating global species richness is to (1)
estimate what proportion of the biota belongs to a particular Current rates of growth
group, (2) estimate what proportion of species in that group
has already been described, and (3) use these estimates and Current rates of description of new species and other taxa
the number of described species in the group to calculate a and how these rates vary from group to group can tell us a
total for all groups. Should the group chosen be the good deal about how the task of inventorying biotic
Coleoptera our estimates might be that this group contains diversity is proceeding. Whether or not rates of description
(say) 20% of all living species, and that (say) only one in have any value for predicting just how much of the task
every five or even ten Coleoptera species has been remains to be done is another matter, considered below.
described. Recent published estimates for the number of Numbers of newly described species recorded in the
described species of Coleoptera, like those for most other Zoological Record for a range of animal groups, for each
large groups, are extremely variable. Several put the figure year between 1979 and 1988, are given in Table 4.5. The
18
Species Inventory
Table 4.3 Numbers of species in the groups of organisms likely to include in excess
of 100,000 species (plus vertebrates)
DESCRIBED ESTIMATED SPECIES
SPECIES HIGHEST FIGURE WORKING FIGURE
Viruses 5,000 500,000+ 500,000 Sy
Bacteria 4,000 3,000,000+ 400,000 Ma/Te/Sy
Fungi 70,000 1,500,000+ 1,000,000 Te/Sy
Protozoans 40,000 100,000+ 200,000 Ma/Te/Sy
Algae 40,000 10,000,000+ 200,000 Ma
Plants 250,000 500,000+ 300,000 Te
(Embryophytes)
Vertebrates 45,000 50,000+ 50,000 Ma/Te
Nematodes 15,000 1,000,000+ 500,000 Ma/Te/Sy
Molluscs 70,000 180,000+ 200,000 Ma/Te
Crustaceans 40,000 150,000+ 150,000 Ma
Arachnids 75,000 1,000,000+ 750,000 Te
Insects 950,000 100,000,000 + 8,000,000 Te
Notes: The figures for described species (mostly given to the nearest 5,000) were arrived at by consulting relevant specialists as well as by critically
reviewing the literature. The ‘highest figure’ estimates for existing species, many of them frankly speculative, are the highest encountered during
a survey of recent literature. The ‘working figure’ estimates are conservative. The figure for bacteria has been arbitrarily ‘capped’ at 100 undescribed
to 1 described species on the grounds that projections involving more than two orders of magnitude are inherently unsafe. The biggest question marks
lie over the true numbers of species of viruses, bacteria and algae. Substantial upward revisions from the working figures for these groups may prove
justified with time. The figures for fungi, protozoans and nematodes are also insecurely based. Note that the Fungi and Protozoa are used in the
‘traditional’ sense, while Bacteria includes cyanobacteria. The figures for ‘insects’ include all hexapods, and that for described insect species assumes
totals of 400,000 for Coleoptera, 150,000 for Lepidoptera, 130,000 for Hymenoptera and 120,000 for Diptera. The final column in the table gives
an indication of where the major proportion of species in each group is concentrated. All groups listed include at least some symbionts (Sy),
obligately associated as parasites, mutualists or commensals with other organisms, and all (except for viruses) have at least some free-living
representatives in marine (Ma), terrestrial (Te) and freshwater systems. Despite high local species richness in some groups, the overall contribution
of freshwater species to group totals is relatively small, unsurprising in view of the fact that freshwater covers well below 1 % of the earth’s surface.
most striking aspect of these figures is the extremely low are as high as or even higher than they have ever been, but
variation between years. Indeed, the yearly overall totals the average figures conceal the fact that the period of
(for all groups included in the Zoological Record) for the maximum species description for a number of groups is
years 1979-1988 show a standard deviation of less than one- well in the past. Detailed information on how taxonomic
twentieth of the mean annual figure. Description rates for activity, as reflected in description rates, has varied through
many of the individual groups listed in Table 4.5 are almost time is available in a range of reviews dealing with
equally invariant through this decade. individual groups, and a summary of this has been provided
by Simon (1983). The picture for groups such as the birds
Description rates can tell us roughly how quickly various is predictable, with most species described early on and half
parts of the taxonomic inventory are growing. As the of the present day total of 9,000 or so recognised species
description of new species in most groups is accompanied having been reached by 1843 (see Fig. 4.1). After a
by continuing reappraisal of the existing inventory, the rate sometimes rather slow start, description of new species in
of description of new species is rarely precisely the same as some groups has otherwise proceeded at a fairly steady rate,
the rate of increase in the number of recognised species. while in others there has been a marked decline after a peak
The description rate may be substantially higher; in some of of activity which in many instances falls towards the end of
the larger groups of insects, for example, the current rate the 19th century. Groups in which the maximum activity is
at which previously described species disappear into taking place now include some of the better-known as well
synonymy is around one-quarter to one-third the rate at as those, such as nematodes and fungi, in which only a
which new species are described. Unfortunately, for some small fraction of species is likely to have been described so
groups, newly recognised synonymies are not systematically far.
reported or recorded in abstracting journals, so that the
extent of the disparity between rates of description and rates Description rates or growth rates do not, of course,
of growth in number of recognised species is extremely necessarily provide a good measure of taxonomic effort
difficult to assess. expended or of how effective this is (see below). A rough
and ready way of determining how such effort is being
How current description rates for a range of groups applied is to look at publication rates (Barnes, 1989, May
compare with earlier rates is indicated in Table 4.6. In 1988). Most instructive, perhaps, is to compare publication
relation to averages for the post-Linnean period, and with rates for various groups with their size, both in terms of
the exception of groups such as birds where few new currently recognised species and also projected overall
species are being discovered, current rates are uniformly species totals. Publication data from the Zoological Record
high. For some groups, such as nematodes, current rates quoted by May (1988) reveal, not surprisingly, that the
19
1. Biological Diversity
Table 4.4 Number of species in various families of beetles (Coleoptera)
DESCRIBED SPECIES ACTUAL SPECIES
Arnett Lawrence All Sources Arnett
(1967) (1982) (1990) (1967)
Byrrhidae 154 c. 300 319 300 TEMP
Derodontidae 10 19 20 19 TEMP
Discolomidae 30 c. 400 443 50 TROP
Dryopidae 178 c. 200 253 300 trop
Elmidae 263 c. 700 1,170 350 trop
Limnichidae 67 c. 200 297 80 trop
Lymexylidae 37 c. 50 64 100 trop
These families in total 739 c. 1,869 2,566 1,195
All Coleoptera 219,409 340,500 ? 290,199
Notes: Number of described species in various families of beetles (Coleoptera) compared with one taxonomist’s estimates (Arnett, 1967) for actual
numbers of existing species in these families. Three sets of figures for numbers of described species are provided. Arnett’s (1985) estimates of
described species were based on catalogues published between 1910 and 1915 while Lawrence’s (1982) estimates were based on more up-to-date
sources. The 1990 figures are based on direct counts from the most recent catalogues available supplemented by Zoological Record entries for
subsequently described species and new synonymies to June 1990. A small part of the increase in numbers of Elmidae recognised in 1990 is due
to species transferred from Dryopidae. If the 7 families for which the 1990 total of described species is 2,566 were representative of the Coleoptera
as a whole in terms of growth since the 1910-1915 period (Arnett’s figures) total described beetle species regarded as valid up-dated to 1990 should
be 761,845 (i.e. 219,409 x 2566/739). Using the more up-to-date estimate (Lawrence, 1982) as starting point the 1990 total for Coleoptera should
be 467,481 (i.e. 340,500 x 2566/1869). In fact, recent species description rates for some of the families in the table (e.g. Elmidae and Limnichidae)
are likely to be well above average for the Coleoptera as a whole, so both extrapolations may produce overestimates. No accurate count has been
made for the whole Order but the actual number of described Coleoptera species regarded as valid as of 1990 is probably in the region of 400,000.
The table illustrates the way in which cautious taxonomists are strongly influenced by the number of species already known when predicting the
number that might actually exist. For relatively well-known beetle groups, largely those in which most species occur in temperate regions, this may
not lead to drastic underestimation, but clearly may do so in the case of less well- known groups in which most species are tropical. TEMP = family
shows a strong bias away from the Tropics; TROP = family shows a strong bias towards the Tropics; trop = family with a weaker tropical bias.
ratio of papers published to number of species already
described has been high in recent years for vertebrates,
varying from around two papers per species in mammals to
one paper for every two or three species in fish. Leaving
aside these well-known groups, the number of papers
published per recognised species per year is more or less
inversely correlated with the size of the group in terms of
described species. For example, in all major groups
including fewer than 50,000 described species the ratio of
publications to species is below 1:50 and often below 1:10,
whereas ratios are generally much higher, exceeding 1:100
in the case of the Coleoptera, in groups containing a greater
number of described species.
Despite this evident bias in taxonomic attention against
groups containing many described species, actual growth
rates, i.e. number of new species described in relation to
the number already described (see Table 4.6), currently
vary remarkably little.
PREDICTION
INVENTORY
FROM THE EXISTING PARTIAL
Inherent limitations
The total number of species so far described, of course,
gives us some idea of the minimum extent of global species
richness, while knowledge of how described species in the
better-known groups are distributed gives us an impression
of the way in which species richness is allocated between
regions, ecosystems, etc. However, there is every indication
that described species do not account for the major portion
of the world’s species and, more importantly, that described
species represent a very biased sample. Is it then possible
to use these figures for described species in any way at all
as a basis for projecting actual world totals?
If we examine the available data on absolute rates of
description and how these have varied through time, and on
how the overall taxonomic effort is and has been
apportioned between the major groups of organisms, we
might justifiably conclude that they tell us very little about
the size of the descriptive task that remains. However, a
careful scrutiny of these data can help to demonstrate the
biases that exist in the inventory as it stands, and may help
to reveal what underlies them. This may be helpful in
making initial judgements as to where the major sources of
unexplored diversity are to be found. It may also prove
useful in assessing the likely accuracy of species richness
estimates, such as those of taxonomic specialists considered
below, that are difficult to evaluate in any other way.
Perhaps the most obvious among biases in how taxonomic
effort is applied are those that stem from everyday human
interests and preoccupations. Large organisms, those that
are considered particularly attractive (flowering plants,
butterflies, etc.) or appealing in some other way, those most
closely resembling humans themselves (vertebrates,
especially mammals), and those that have a direct impact on
human affairs, usually as pests of one kind or another, are
all favoured objects of study and description. By
association, certain other groups such as fleas and lice -
because they are parasites of birds and mammals - may also
receive a relatively large share of attention.
Species Inventory
Table 4.5 Number of new species listed in the Zoo/ogical Record 1979-1988
1979 1980 1981 1982
Protozoa 463 309 353 280
Platyhelminthes 293 357 341 290
Nematoda 407 389 364 422
Annelida 234 114 215 186
Mollusca 338 355 391 239
‘Other’ invertebrates 180 214 341 174
Crustacea 645 638 763 736
Arachnida 1;oJOpe 523 1,199 1,182
‘Other’ arthropods 98 57 120 154
Hemiptera 1,277 1,115 1,191 981
Lepidoptera 675 445 506 580
Diptera 1,019 1,015 1,130 899
Hymenoptera 1,167 1,134 1,086 1,031
Coleoptera 2,116 2,804 2,243 2,454
Other Insecta 725 824 992 878
Pisces 183 241 273 240
Other Chordata 146 170 134 186
1983 1984 1985 1986 1987 1988
272 374 432 385 315 377
307 340 359 336 297 243
383 354 365 367 331 258
127 208 200 149 161 131
344 316 419 412 392 419
303 353 187 252 195 339
695 599 700 843 708 660
il45> 15047 is53" 1/185) 1488) 1307
119 123 55 94 146 122
1,275 1,038 1,072) 11162 903 1,016
631 685 658 915 623 699
973 928 1,115 1,095 1,303 1,000
1,215 1,496 857 1,184 1,084 1,705
1,960 2,259 2,220 2,843 2,130 2,051
1,331 1,030 836 822 1,110 703
260 223 220 204 234 229
177 168 230 117 191 138
Source: Zoological Record Online, data search organised and carried out by BIOSIS UK.
Notes: Number of new species listed in the Zoological Record 1979 to 1988 (Vols 116-125), showing the remarkable constancy of description rates
in the larger animal groups. The figures given are for newly described species. While extinct groups such as the Trilobita are excluded, fossil species
(mostly relatively few in number) of extant groups (e.g. Mollusca, Insecta, Pisces) are included in the counts. As new synonymies are not accounted
for the figures do not provide a precise measure of growth in numbers of described species regarded as valid. The ‘other’ invertebrates category
includes all extant non- arthropod groups not otherwise listed. The ‘other’ arthropod category includes all taxa listed in Section 12 of the Zoological
Record; in major part these are Chilopoda (centipedes) and Diplopoda (millipedes). The other Chordates category includes reptiles, mammals,
amphibia and birds.
Other strong biases have more to do with taxonomic taste
and fashion, and the ease with which the organisms are
found, collected, studied and preserved as specimens.
Organisms that can be studied without complex procedures
or expensive equipment, that are not too small, that exhibit
distinctive characteristics and can be readily sorted tend to
be dealt with preferentially. These various biases have
influenced the historical pattern of description, starting with
Linnaeus himself, who described very few small organisms,
and confirmed by Gaston (1991b) who showed a clear
relationship between body size and date of description in the
approximately 4,000 species of British beetles.
It is also evident that most areas distant from the centres of
human population as well as more obviously inaccessible
regions such as the ocean depths are rather poorly
inventoried. It should also be noted that most taxonomists,
however much they may travel, remain based in the cities
of the north temperate zone. Recent patterns of description
for birds and mammals suggest that the few species that
remain to be discovered in such well-known groups will
almost certainly turn out to be tropical. Despite clear
indications that the greater part of global species richness is
to be found in the tropics, it is evident (no precise counts
are available) that, in terms of described species, those of
tropical origin are considerably outnumbered by those from
temperate and boreal regions. An analysis of recent
description data for insects (Gaston, unpublished) reveals
that, for this speciose group at least, the bias against
tropical species is still not being redressed. In some groups,
the rate at which new species are described may be
determined, at least in part, by what material is immediately
available for study, which in turn depends on its general
accessibility in nature. However, this can scarcely be the
major influence on description rates in those groups (e.g.
the larger insect orders) for which the world’s museums
already contain hundreds of thousands of undescribed
species.
Time-series of species descriptions
Changes in the rate at which new species have been and are
being described have been used to make an estimation of
the likely future growth of each of the major groups of
organisms. Various statistical procedures can be used to
identify trends and project description rates forwards (May,
1990), with different techniques sometimes producing very
different results (Frank and Curtis, 1979; Simon, 1983).
Not surprisingly perhaps, attempts to use trends in
description rates to predict global species richness of major
contributors (i.e. excluding groups such as birds, mammals,
etc.) or of the biota as a whole have been singularly
unsuccessful (see Erwin, 1991). First of all, as a glance at
Table 4.5 reveals, very few trends in description rates can
be observed over the short term. Longer term trends, where
they are evident, are often erratic but, except for a few
small groups, commonly involve an increase in the pace of
description over time. Species description in the least
apparent and least tractable groups often gets off to a slow
start but once this gets going, description rates may be
more or less monotonic.
For those few groups in which species description is
1. Biological Diversity
Table 4.6 Current species description rates for various animal groups and for fungi
SPECIES ‘GROWTH’ CURRENT PROPORTION
DESCRIBED RATE RATE/ OF SPECIES
PER ANNUM PER ANNUM OVERALL DESCRIBED
(1978-1987) (1978-1987) RATE TO DATE
Vertebrates 367 0.82 1.90 High
Birds 5) 0.05 0.13 Very high
Mammals 26 0.59 1.37 M
Amphibians and
Reptiles 105 1.17 222 High
Fish 231 1.22 2.83 z
Molluscs 366 0.52 qe22. Moderate
Sponges 50 0.56 1.30 y
Cnidarians 57 0.63 1.48 *
Platyhelminths 316 1.58 3.68 yy
Ectoprocts 58 ied) 3.00 .
Annelids 173 115 2.57 "
Protozoans 356 0.88 2.00 Moderate/low
Crustaceans 699 1.74 3.91
Insects 7,222 0.76 1.77 Low
Lepidoptera 642 0.43 1.00 Moderate/high
Coleoptera 2,308 0.57 1.34 Low
Diptera 1,048 0.87 2.03 Low/very low
Hymenoptera 1,196 0.92 2.14
Arachnids 1,350 1.80 4.19 es
Fungi 1,700 2.43 5.67 Very low
Nematodes 364 2.43 5.65
Notes: Current species description rates for various animal groups and for fungi, expressed as number of species described per annum (mean of
years 1978-1987) (Column 1), compared with number of already described species (figures from column 1 (x 100) divided by number of described
species) (Column 2), and with an approximation to average description rates for the whole of the period since 1758 (mean number of species
described per annum 1978-1987 divided by mean number of species currently recognised as valid described per annum between 1758 and the present)
(Column 3). The fourth column provides an indication of the proportion of each group that is likely to have been described so far; very high = c.
90% or more already described; high = c. 50-90%; moderate = c. 20-50%; low = c. 10-20%; very low = less than 10%
nearing completion, description rates may be expected to
have some predictive value. Even here, however, they are
likely to tell us what we already know, and may be
distinctly misleading. Growth curves based on time-series
of species descriptions are a convenient way of portraying
the relevant data. The time-series for the very well-known
groups such as birds generally forms a classic S-shaped
growth curve (see Fig. 4.1, but note that here the curve is
not S-shaped as the vertical axis is on a logarithmic scale).
Any other form of curve indicates that the group in question
is unlikely to be almost completely inventoried, but may tell
us little else. We may note that birds are exceptional among
relatively high-ranking taxa in that the description of new
species has slowed to a trickle. Even mammal species (see
Table 4.6) are still being described at a rather high rate that
gives no clear sign of an asymptote, although it is fairly
certain that the number of species awaiting discovery is
relatively low. Some slowing down of the rate, finally, is
perhaps indicated by figures for the last decade or so, with
an average of 37 mammal species described per year from
1978 to 1982 and 20.5 per year from 1983 to 1988.
There are various ways of gathering the data for time-series
graphs, and these can have significant effects on our ability
to predict. For example, the number of species recognised
within a group at any one time can be assessed from
22
contemporary taxonomic works. However, over the past
120 years, there have been major changes in how species
status is evaluated, which can make the figures
non-comparable and the curves uncertain.
A better curve is usually obtained by making a cumulative
graph of the dates of first description of all currently
recognised species. In the case of the 120 or so species of
crows (Fig. 4.2A), for example, we see a truncated version
of the S-shaped curve, with nearly 10% of all currently
recognised crow species having been described within the
decade 1758-1767. With the curve for crows flat for the last
quarter century it would be a bold person who would
predict a rise to even 130 species, let alone a higher figure.
However, such dramatic shifts can occur, even in relatively
well-known groups.
An example is provided by a group of blue butterflies (Fig.
4.2C), subject of a recent major revision (Eliot and
Kawazoe, 1983). Linnaeus knew only one species, the
familiar European Holly Blue Celastrina argiolus, and the
time-series is very slow up until the decade ending 1877.
After that it goes through a rapid growth-phase, and then
flattens at about the same time as the curve for the crows.
Since 1967, however, there has been a new burst of species
description, bringing the current total to a point at least
Figure 4.1 Discovery curves for species
from 1758 to 1970
125 years
log scale
0.01
178 1843
1970
Arachnids and Crustaceans
10
0.5
202 years
log scale
0,001
1960 1970
1758
Source: Following May (1990) after Simon (1983).
Notes: Numbers of known species (expressed as a fraction of those
known in 1970 on a logarithmic scale) are plotted against time. The
vertical and horizontal lines show the points at which half of the 1970
totals had been reached. Although a trickle of new species of birds
continues to be described the shape of the curve for birds as a whole
resembles that for crows. The curve for Arachnida + Crustacea (i.e.
the majority of non-insect arthropods) shows that up to 1970
description of new species had an ever-increasing pace, with the 1960
total doubled by 1970. Description of new species in these groups now
proceeds at a steady rate of some 2,000 per annum (see Table 4.5).
33% higher than the plateau level. The explanation here is
not poor taxonomy or a change in species concept, but a
combination of exceptionally painstaking work coupled with
vigorous collecting in previously inaccessible parts of
Southeast Asia, where these butterflies form many island or
mountain endemics.
23
Species Inventory
OTHER APPROACHES TO PREDICTING PATTERNS
Estimates by taxonomic specialists
The opinions of taxonomists specialising in particular
groups of organisms have traditionally played a considerable
part in the formulation of views on the extent and pattern of
species richness at every scale. Indeed, the preliminary
tentative working figures for global species richness of the
major groups used in this section have inevitably been
influenced by the opinions and estimates of relevant
taxonomists. However, the simple approach of collating
views based on the specialist knowledge of the taxonomic
community has not been systematically pursued, a major
exception being the recent essay by Gaston (1991a) to
assemble and interpret a cross-section of taxonomists’
opinions concerning likely global insect species richness.
The approach adopted by Gaston has the merit of involving
a large number of data points so that no one estimate has an
overriding effect on the overall result. In addition, the
sources are experienced taxonomists whose work generally
involves exposure to at least part of the richness of species
located in poorly studied regions. This said, it is likely that
the way in which taxonomists actually arrive at their
conclusions is quite varied, may be distinctly idiosyncratic
and tends to the conservative. Indeed, the generally rather
poor track record for such estimates suggests a possible
correlation between the degree to which any given
taxonomist has been exposed to relevant data (e.g.
representative samples from many areas, including some of
the richest) and the extent to which he or she is prepared to
extrapolate beyond the relatively sure ground of already
described species. Very early estimates by such as John Ray
who, in the late 17th century, considered that the insects of
the world as a whole might amount to some 10,000-20,000
species, may lend some support to this view.
To the extent that taxonomists work largely with what
happens to come their way, it is likely that the collections
they examine do not fully represent the richness to be found
in less-known regions of the world, such as the tropics. In
making their assessments of overall species richness it is
also likely that they make some use, however
unsystematically, of described to undescribed species ratios
(see also below) in the small groups with which they are
most familiar. If the group already contains (say) 100
nominal species, and the taxonomist in question is aware
that 10 of these are not ‘good’ but is also aware of a further
60 undescribed species, the new provisional total for the
group will be 150 species, representing an increase of 50%.
The value of this figure for generalising will, of course,
depend very much on how typical the sample group is and
how well the available material represents its true size.
Nevertheless, if accepted for what they are, and if we
accept also that recent estimates by taxonomists are based,
in comparison with their predecessors, on a relatively
extensive (if still fragmentary) coverage of the world, the
surely conservative figures produced by the cautious and
pragmatic approach may have considerable value as
minimum estimates.
Gaston’s conclusions have attracted strong criticism (Erwin,
1. Biological Diversity
1991), the main focus of which is that the reliability of
results obtained in this way is impossible to judge;
taxonomists’ estimates represent opinions that have been
arrived at in ways that we cannot know. The arguments for
and against have broadened to include the merits of other
approaches as well as the usefulness of collated opinion,
providing an area of active debate (see Gaston, 1992).
First principles and empirical relationships
The broad understanding we have of how life evolved and
how species interact could be used to estimate, from first
principles, how many species are likely to be found in a
given region or in the world as a whole (May, 1988).
General rules concerning: body size relations, commonness
and rarity, range sizes, and the relationship between species
numbers and area have all been used to suggest explanations
for observed species richness patterns and why there are so
many (or so few) species overall. Understandably, only
tentative use has been made of rules of this type for actually
predicting major species richness patterns for poorly-known
groups. Any real test of their predictive power in these
areas awaits the provision of many more data concerning
the exceptionally diverse but little-known groups than are
available at the moment. This applies, for example, to the
empirical rules, derived mainly from the larger terrestrial
animals, that describe the way in which species numbers
increase with decreasing size. Using only described species
these rules begin to break down at body lengths of below
about lem. Arbitrary extrapolation to smaller size classes
(down to lengths of about 0.2mm) that are poorly
represented among described species produces an estimated
global total for terrestrial animals of around 10 million
species (May, 1988).
The use that may be made of other empirical relations that
concern the structure of food webs, and the numbers of
parasitic or other symbiotic species that are typically
associated with individual host species, has also been well
reviewed by May (1988, 1990). While rules concerning the
number of levels in food webs are sufficiently well
established to form the basis for relatively reliable
generalisation, the same cannot be said for the numbers of
species and overall numbers of links involved in webs of
various types. Species richness patterns involving parasite,
parasitoid or (less often) predator species and their hosts or
prey have received much attention. In well-known regions
such as the British Isles it is possible to calculate the
approximate number of potential host species for a given
group of, for example, parasites and relate this to the
overall number of species of these parasites that are present.
If we take British vascular plants (2,089 species) and the
insects that directly exploit them (assuming this to be
around 25% of the British total or c. 5,500 species) as an
example, we can derive a ratio, in this case of around 2.6
(associated insect species) to 1 (plant species). This type of
simple relationship tells us very little, of course, about
host-specificity. Nevertheless, the question of host-
specificity levels, rather than any empirical relationship
between the number of hosts and the number of associated
parasites, has received some attention as a possible means
of predicting overall numbers of parasite species. The
difficulties involved in evaluating such patchy
host-specificity data as exist and using them for
extrapolative purposes are great (see May, 1990 for
discussion).
Keeping to vascular plants and their associates as the
example, we see that the most useful data on how many
species may be effectively specialised to one host come
from detailed single species studies. Intensive studies,
whether of oak trees or passion vines (see May, 1990), may
help to reveal something of the processes underlying the
way in which these plants are exploited, while at the same
time elucidating a series of contrasting patterns. However,
they cannot be expected to provide what is required for any
prediction of overall numbers of plant associated species.
The simple questions for which answers are needed here
are: how many species depend on the average plant species
throughout its range, and how many species depend on the
same average plant species in one place at one time? For
practical purposes it is also advantageous if these data can
be related to sampling phenomena, so that it is known what
proportion of the associated species present at one place are
obtained in a particular type of sample.
As far as species associated with green plants are concerned
there are indications that patterns vary with moisture,
latitudinal and other gradients. Host-specificity levels may
also tend to be lower where plant species richness is
especially high, particularly when the plants in question are
trees, as in tropical moist forests. Indeed, there are strong
suggestions that the general architecture of forests may be
a better predictor of the number of small animal species and
fungi present in a given area than is the number of different
vascular plant species that occur.
Taxon to taxon and region to region relationships
Using some aspect or aspects of the diversity profile of a
well-known group such as birds or mammals as a reference
point, a variety of simple extrapolations to other less
well-known groups may be made. We may use butterflies,
a well-known group, as an example. Of the roughly 22,000
species of insects to be found in Britain some 67 are
butterflies. The number of described species of butterflies
in the world is fairly accurately known at around 17,500,
the true figure almost certainly not exceeding 20,000 or so.
If the ratio of butterfly species to all insect species is the
same globally as it is in Britain then the world insect
species total should lie at around 22,000 x 17,500/67, that
is 5.75 million.
A more involved extrapolation may be made by taking
tropical to extratropical ratios as the point of departure. For
both birds and mammals, for example, there are roughly
two to three times as many tropical as non-tropical species.
To extrapolate successfully from this we need to have a
good estimate of the proportion of described species that are
from extra-tropical areas in the more significant of the less
well-known groups, coupled with a good estimate as to the
proportion of extra-tropical species that have been
described. In practice our estimates for the first are unlikely
to be very accurate and for the second unreliable. However,
Species Inventory
Figure 4.2 Time series of first descriptions of currently recognised species in
decades from the time of Linnaeus (1758) to 1987
100
Number of Species
so
Danainae (160 species)
Lycaenopsis (112 species)
1757. (1767 «#1777 #1787 #1797 #1807 1817 1827 1837 1847 #1857 1667 1877 1687 1897 1907 1917 1927 1937 1947 1957 1967 1977 19867
Time
Source: Data for crows (Corvidae) based on Goodwin, 1986, Crows of the World, London: BM(NH), that for milkweed butterflies (Danainae) on
Ackery and Vane-Wright, 1984, Milkweed Butterflies, London: BM(NH), and that for the Lycaenopsis group of blue butterflies (Lycaenidae on Eliot
and Kawazoe, 1983, Blue Butterflies of the Lycaenopsis Group. London: BM(NH).
Notes: Asterisks on each curve indicate the points at which half of the 1987 totals had been reached. All three of the groups depicted are
‘well-known’ with few if any species left to be discovered and described. The rate at which new species of crows (Corvidae) were recognised and
described declined steadily from the mid 1800s so that 90% were known by around 1880. Description of milkweed butterflies (Danainae) followed
a largely similar pattern, with 90% of the apparently settled total achieved by 1937 and maintained for the next three decades also reached by around
1880. However, intensive studies over the past two decades have led to a further (and unpredicted) small burst of description. After a much slower
start, the Lycaenopsis group of blue butterflies (Lycaenopsis) also reached a seemingly stable plateau (by around 1920). As with the milkweeds an
unpredicted burst of description, although in this instance a much larger one, has characterised the last decade or so.
again taking insects as the example, if we take one million
as the rough number of described species, and assume that
(1) roughly 60% of described insect species are from
temperate and boreal regions, and (2) 40% of extra-tropical
species have been described, then ratios of two or three
tropical species to one extratropical species give us world
insect species totals in the range 4.5-6 million. There are
few suitable data points to use for microorganisms and
some of the other groups such as nematodes and mites
discussed below, even for north temperate sites and regions,
but extrapolations based on the pattern of species richness
in vascular plants, various vertebrate groups and on
butterflies all produce roughly the same kinds of answers
for the remainder of the biota, including the insects.
All such calculations, of course, depend on how similar
bird, mammal, butterfly or other patterns used in
calculations are to those found in the much richer but less
well-known groups. If we were, in fact, confident that
patterns found in groups such as birds were universal we
would be close to achieving reasonable understanding of the
global picture. But just how ‘typical’ are these well-studied
groups with respect to species richness patterns, including
their local species richness in tropical as opposed to
temperate areas, and the rates at which species accumulate
as the area considered is enlarged? We know enough to be
clear that latitudinal gradients of species richness are not the
same in all major groups (although species richness does
generally increase dramatically with reducing latitude).
Turnover rates also vary substantially from group to group,
although evidence presently available (mostly of course for
well-known groups) fails to reveal any clear correlation
between these rates and size or other significant biological
attributes that might suggest large average differences
between (say) mammals and small invertebrates. A more
rapid turnover in tropical as opposed to temperate regions
does, however, seem to be indicated by the evidence, and
various explanations for this have been advanced.
However, in the absence of data that might be used for
more direct approaches to calculating species richness in the
largest and most poorly-known groups, simple
extrapolations from well-known groups are likely to provide
us with the most securely based, if very conservative,
estimates attainable at present. To do distinctly better it will
be necessary to identify clearly which of the poorly-known
1. Biological Diversity
groups might eventually make a major contribution to the
taxonomic inventory (see below), and gather fresh relevant
data by direct sampling from nature.
The relationship between the number of described and
undescribed species in any group requires comment. The
usefulness of this relationship as a means of predicting the
number of species in a group depends on the extent to
which representative samples are available and the accuracy
with which the proportion of species that are undescribed
can be ascertained. In practice, the latter is generally time-
consuming and difficult, if not impossible. Unfortunately,
where most feasible (e.g. in very small groups and groups
in which most species have already been described), the
results obtained will tend to be uninformative. Where the
approach is potentially most valuable (e.g. very speciose
groups in which 75% or more of the species remain
undescribed), it is most difficult to apply. Here, there is a
premium on accuracy, but this can only be achieved by
someone who has close familiarity with all of the described
species that might be present in the sample. Nevertheless,
the effort may be worth making for groups likely to make
a major contribution to global species richness. Any
indication as to whether undescribed species are, for
example, around three times as numerous (i.e. 75%
undescribed) or (say) 19 times (i.e. 95% undescribed) as
numerous as described species would be of considerable
value.
UNCHARTED REALMS OF SPECIES RICHNESS
Here we turn away from the existing taxonomic inventory
and knowledge of species richness patterns in well-known
groups to consider directly where the major part of as yet
unassessed species richness might lie. For which
ecosystems, taxonomic or other groups are there indications
of great unassessed species richness? Is it possible to
pinpoint the areas that it is essential to take into account if
global totals are to be roughly estimated? Included in the
discussion are the principal among the biological ‘new
frontiers’ that have attracted attention in recent years.
Evidence or the presumption that local species richness is
at least sometimes high provides the first hint that a
taxonomic group or a type of community might make a
large contribution to the global species total. However, in
sifting the stronger indications out from less telling
anecdotes or the merely hyperbolic, it is helpful to
remember that high local species richness, although
necessary, by no means provides a sufficient demonstration
that the group in question makes a particularly large
contribution overall.
The marine realm
The oceans, occupying over two-thirds of the Earth’s
surface, have been described by Colinvaux (1980) as
making up "a vast desert, desperately short of nutrients and
with living things spread most thinly through them". This
blunt description, dismal as it may seem, nonetheless
provides an effective summary of what is known of marine
productivity, turnover time and biomass. Average biomass
(per unit area) in the seas has been estimated to be of the
order of one thousandth that on dry land while marine
productivity (again per unit area) is about one-fifth of the
26
average for terrestrial systems (Valiela, 1984). In absolute
terms it has been calculated, for example, that the world’s
seas produce some 92,000 million tons of plant tissue per
annum, as against 272,000 million tons for dry land plants.
Although new data may necessitate some revision of figures
of this type they are unlikely to change the general picture.
Against this background it may be unsurprising that there
are few data to suggest that the oceans contribute more than
a small fraction to the world total of species, at least of
multicellular animals and plants. In contrast, the marine
realm makes an exceptional contribution to biotic diversity
at higher levels (all major eukaryote groups are represented
and more than 80% of all phyla are restricted to the seas).
Of all currently described species it has been estimated that
somewhat less than 15% are marine. The views of relevant
taxonomists (see Barnes, 1989, etc.), supported by the
generally rather high proportion of described species in
samples taken from poorly studied areas, suggest that fairly
high percentages of the marine ‘macrofauna’ (mostly
species of molluscs, crustaceans and polychaete worms) and
multicellular algae are already known. The position with
regard to smaller organisms, including nematodes and
protists, is very much less certain. Moderately high species
richness at the local level can be found in some inshore
communities where productivity is high, those of tropical
reef systems providing good and well documented
examples. However, total areas occupied by these rich
communities are small and many of the species have fairly
large ranges; thus local species richness of the apparently
relatively well-described littoral and shallow water marine
communities is not reflected in especially high regional or
global described species totals.
Although the ranges occupied by most marine organisms are
poorly understood, patterns observed in the better-known
groups suggest that turnover of species, the rate at which
species numbers increase with increasing area, may be
generally lower in the seas, perhaps especially in the open
oceans and the ocean depths. Unlike the continents the
oceans are contiguous; also the deep sea appears to have
few areas sufficiently isolated for boundaries to be defined
and thus few limits to dispersal which, even for small
sediment-dwelling animals, may be through planktonic
larvae. Although volumetrically great, the seas are also
architecturally not very varied. As noted above, systematists
working on most marine groups (see Barnes, 1989) appear
reluctant to suggest that large numbers remain to be
described and, compared to terrestrial arthropods, for
example, this may well be true for such groups as
Echinodermata, the larger Mollusca and Crustacea, etc., as
well as fishes.
The deep sea is one of the more remarkable biological ‘new
frontiers’ that has become evident in the past few decades
(see Grassle, 1989, 1991; Grassle er al., 1991). Although
some parts of the deep sea floor are apparently poor in
species, high local species richness of macrofauna in deep
sea sediments appears to be the rule over the fairly large
areas that have now been investigated in the Gulf of
Mexico, the West Atlantic (Grassle, 1991) and elsewhere.
This is manifest mostly among polychaete annelids, certain
groups of Crustacea and, to a lesser extent, molluscs. Low
productivity, sediment patchiness and ease of immigration
are among the factors suggested to explain this diversity.
Distinct depth and sediment type assemblages have also
been shown to occur,. but there is little indication in the
macrofauna of high turnover across all spatial scales.
Indeed, the major part of local species richness seems to be
exhibited at a very small scale, so that the majority of
species to be found at one site are obtained by very few
samples. The smaller organisms or meiofauna of deep ocean
sediments often equal the macrofauna in biomass and are
present in much greater abundance, the major component
being nematodes. However, whether meiofaunal species
richness equals or possibly exceeds that of the macrofauna
remains to be established. Although relevant data may be
forthcoming from studies in progress, as yet how nematode
species of deep ocean sediments accumulate as we move
from site to site is more or less unknown.
Although these new data on the deep sea, coupled with
recent discoveries of a whole new realm of protistan,
bacterial and other picoplankton suggest that total marine
biotic diversity could be considerably greater than
previously assumed, evidence to support the contention that
this richness rivals that found in tropical forests, except
perhaps at the smallest of scales (i.e. the range below 1m?)
is wanting. New data on both pelagic and benthic
microorganisms and the deep sea meiofauna may yet
confound this view, but the evidence so far suggests that the
oceans, including their poorly explored depths, contribute
less to total global species richness, by an order of
magnitude or more, than do moist tropical forests.
Parasites
Parasite loads for a few large animals (mostly vertebrates)
and some green plants may be high, involving many
parasites that are specific to a single host or a narrow range
of host species. However, the overall numbers of large
animal and large vascular plant symbionts, unless there are
many more unknown than we suppose, are insufficient in
themselves to make a very large contribution to global
species richness. In contrast, very little is known
concerning loads and levels of host-specificity with respect
to the microorganisms, small nematodes, mites and others
that are associated as parasites with members of the most
species-rich groups, such as terrestrial arthropods. In
relatively well-known areas such as the British Isles the
recorded numbers of such parasites are low, but even here
it is not unusual for small invertebrate animals to turn out
on close examination to possess previously unknown
parasites. Clearly, if there are many such undetected
parasite species, their numbers could lead to a considerable
inflation of global species figures. For example, if each
insect species has, on average, one completely specific
associated parasite or other symbiont this would entail at
least doubling estimates of insect species to obtain a
minimum figure for overall global species richness. As yet
there is little evidence that this may be necessary, as where
a range of insects and other small potential host species
have been relatively well studied, large numbers of
host-specific parasites have not been found. We may note
that such negative results (absence of parasites) often go
unremarked and unreported. There is also an inevitable
general tendency for host ranges to be underestimated. In
addition, it is reasonable to assume that the sometimes high
27,
Species Inventory
parasite loads observed in widely distributed pest species
are not, in fact, typical, and furnish a poor basis for
extrapolation. We should also not be too eager to generalise
from the situation in large vertebrates and vascular plants
whose size and bodily complexity furnish many potential
niches for exploitation. The great majority of organisms,
small in size, clearly offer very different opportunities to
potential parasites. On first principles, levels of parasitism
may be expected to vary very widely, depending not only
on the size of the host but also its defences and its
population structure. Potential hosts that are very hard to
find will generally have few obligate parasites.
Fungi and microorganisms
Although far fewer species have been described than of
green plants it has long been considered likely that the fungi
(using the term in its traditional non-phylogenetic sense)
might eventually prove to be the most species-rich of all
groups, insects excepted. Interestingly, at a time when only
a few thousand species of fungi had been described, some
19th century mycologists early on recognised the likelihood
that some hundreds of thousands might actually exist.
However, with around 70,000 described species now
recognised, we are still not in a position to say much more
than this about the size of the group. In the absence of good
data on tropical fungal communities, on latitudinal or other
gradients in diversity, and how the numbers of fungus
species accumulate as we move from one spatial scale to
another, any estimates of overall fungus species richness
can only be tentative.
In a thorough review of the significance and possible
magnitude of fungal diversity, Hawksworth (1991, and see
this report) has settled on 1.5 million as a conservative
estimate for the world’s species of fungi. This figure was
arrived at by taking into account several types of evidence,
but finds its most firm basis in the relationship between the
number of species of fungi known to occur in the British
Isles and the number of British species of vascular plants.
The list of fungus species recorded from the British Isles
currently stands at around 12,000. Taking a figure of 2,089
(i.e. garden species, etc. excluded) for British vascular
plant species, we arrive at an approximately 6:1 ratio in
favour of the fungi. Applying this ratio to a conservative
global figure for vascular plant species of 270,000 yields a
global total for fungi of around 1.6 million species.
As already discussed above, the reliability of extrapolations
made in this way depends on the extent to which species
richness patterns are shared, in this instance between fungi
and vascular plants. At least some fungus species have
extremely large ranges; should average range size in fungi
be significantly greater than the average in vascular plants,
some lowering of the 1.5 million figure for fungi would be
in order. Similarly, should fungi exhibit a less steep
latitudinal gradient in species richness than that found in
vascular plants, this should also point to a lower figure.
Data on tropical fungi remain extremely scant, but we may
note that the rather low proportions of undescribed species
found in recent tropical collections as yet provide no
indication of especially great tropical diversity.
Taking a cautious approach similar to that adopted here
1. Biological Diversity
towards other poorly-known groups, a minimum figure for
global fungus species might be put at around half a million.
An alternative, less cautious but well-supported, approach
is presented in Chapter 6 of this book. The arbitrary
‘working figure’ of one million incorporated in Table 4.3
represents a compromise between this and the 1.5 million
estimate given by Hawksworth (1991).
Microorganisms, including the smaller fungi, algae and
‘protozoans’, as well as bacteria and viruses, present the
greatest challenge to any serious attempt to assess the
overall scale of global species richness. The great genetic
diversity and general significance of microorganisms is
highlighted in Chapter 6, where the problem of applying to
them the species concepts that are more or less consistently
used for many larger organisms is also discussed.
What is clearly an immense diversity of very small
organisms, perhaps especially bacteria, viruses and
unicellular algae, remains largely unaccounted for by the
existing taxonomic inventory. However, whether the
diversity of these organisms, often lacking sexual processes
and many of them clonal, is best expressed in terms of the
number of phenetic groups recognised as species is a moot
point. The comparability of, for example, viral ‘species’
and those of multi-cellular organisms, in which sexual
reproduction predominates, is very questionable. Virtually
nothing is known of any latitudinal or other gradients of
diversity that microorganisms might exhibit while, even in
temperate regions, at no scale is species richness well
documented. Probable range sizes are also known for very
few species, but very small organisms (and those with very
small dispersal stages, such as fungal spores) are known, in
some instances, to have very broad if not cosmopolitan
distributions. Coupled with a generous measure of caution
in extrapolating too far from the known, all of these
considerations are reflected in the arbitrary ‘working
figures’ for species richness of microorganism groups given
in Table 4.3.
Nematodes, mites and insects
Despite a considerable increase in resources devoted to
nematode taxonomy over the past few decades and a
commensurate surge in the rate of description of new
nematode taxa, this group of worms probably still remains
the least well inventoried group of metazoan animals.
Although relatively early attention had been devoted to
some of the larger and, in human terms, more significant
parasitic species, up until 1860 only 80 species of plant,
soil and freshwater species had been described. This
compares with an annual rate of around 140 species of the
same groups described in the 1960s and the present overall
description rate (including parasitic and marine taxa) of
more than 300 species per annum. The current total of
described species is very uncertain but has been estimated
to stand at around 15,000.
Nematodes
Indications that nematode species richness may be of an
extremely high order stem more than anything from the
abundance of free-living forms (a few millions of
individuals may be present in 1km? of suitable soil or mud)
and the great number of free-living species that may be
found in samples taken from a very small area. Two
hundred or more species have been reported from samples
of just a few cm? of coastal mud.
While parasitic species totals may prove to be significantly
high (see above), and free-living terrestrial and freshwater
species also very numerous (Poinar, 1983), recent work on
estuarine, shallow-water and deep-sea sediment nematodes
suggests that the marine realm (see above) could make an
even greater contribution to a total count of the world’s
nematodes. However, how high levels of species richness
at the smallest scales bear on the question of the overall
number of nematode species remains unclear. Good data on
species turnover in both terrestrial and marine nematode
assemblages are conspicuously lacking, as is any indication
that assemblages of tropical nematodes are especially rich.
In the absence of any direct indication of massive
unaccounted for species richness at larger scales a
somewhat cautious approach to estimating the likely overall
number of nematode species is probably advisable.
However, it would be surprising if this number were not at
least some hundreds of thousands.
As in the case of protists and other microorganisms the
taxonomic study of nematodes is made difficult by
uncertainties with regard to the application of species
concepts. Many species are entirely uniparental or contain
some uniparental populations. Apart from their frequently
very small size, the sorting to species of nematode samples
is often hampered by a very low incidence of diagnostic
males. At best, species recognition is beset by many
difficulties and may, in some instances, remain frankly
subjective.
Mites
In the case of mites (Acari) there are fewer problems with
interpreting species limits but, as with nematodes, the
number (around 30,000 or so) of described species clearly
represents only a small proportion of the actual total.
Knowledge of tropical mite faunas in particular is very
scant, lagging well behind that of other arachnids, including
spiders. Reliable quantitative sample data that give anything
more than a hint of what mite species richness might be at
any site in the tropics appear to be unavailable. However,
it may be reasonable to expect that free-living terrestrial
mites, although flightless and differing from insects in
various other respects, do roughly follow patterns, in terms
of coexistence, range sizes, turnover, etc., already
tentatively established for certain insect groups. If so, and
despite the fact that we have a less complete knowledge of
temperate mites than, say, of beetles, it is difficult to
envisage a world total of less than a few hundred thousand
species. Suggestions that the global number of mite species
is in the region of one million or even higher may prove
defensible once good data for tropical sites are forthcoming.
Insects
There is abundant evidence to suggest that insects exhibit
high species richness at most scales (i.e. from a few m* to
ecosystems) except perhaps the very smallest. The number
of already recognised and described species - around one
million - is sufficient to establish that insects comprise a
substantial portion of the world’s species. Most insect
groups are taxonomically tractable and the rate at which the
process of inventorying advances depends largely on the
level of resources devoted to the task. Samples containing
many species can often be fairly rapidly as well as reliably
sorted, and this makes several major insect groups suitable
for a range of species richness studies, even when most of
the species being examined are undescribed. Some of the
ways in which data from samples of tropical insects may be
used to tackle the problem of assessing insect global species
richness are discussed below. We may note, however, that
attaining any reasonably accurate idea of what proportion of
species in total are insects is less easy. This is likely to
depend as much on achieving advances in estimating the
diversity of microorganisms and other poorly understood
groups as on better data for the insects themselves.
Tropical forest canopies: the height of tropical diversity?
Tropical forests have long been known to harbour a great
richness of life and, although they cover only 6% of the
earth’s land surface, it has been widely supposed that they
may contain as many species of organisms as, or even more
than, the rest of the world together. One part of these
forests, the world of the tree tops, has tended to evade close
inspection by biologists but, with the development over the
past two decades of new methods for studying forest canopy
organisms, notably (but not only) the use of insecticide
fogging techniques, canopy communities even in tall
tropical forests have become much more accessible (Erwin,
1990). -
There is now sufficient information to indicate that local
species richness of many of the insect and other arthropod
groups that have been the main focus of recent attention are
very high in tropical forest canopies, much higher (often by
a factor of 10 or more at the level of a single tree) than in
temperate forests. It is equally clear that not only are a high
proportion of the species undescribed (this is the case for all
strata in moist tropical forests) but a proportion of them are
not or are only exceptionally found at lower levels. Data
have now been gathered that give some idea of the usual
sort of numbers of species of at least some of the more
important insect groups (notably Coleoptera and Hemiptera)
to be found in various neotropical and palaeotropical forest
canopies at the level of individual trees and small quadrats
(e.g. 12 x 12m), up to about the one hectare level.
Fewer data are available to allow confident estimation of
canopy species numbers at a larger scale within relatively
uniform tropical forest. Indications are that much of the
patchiness in the canopy is at or below the one hectare level
and that samples from adjacent hectares are about as
different in species composition as samples taken several
kilometres apart. The picture that is beginning to emerge is
of a mosaic less defined by tree species than by a variety of
other factors, including the condition of each tree, and the
patchwork distribution of resources, including epiphytes,
that manifests itself at a much smaller scale than an
individual tree canopy. Some data are available to show that
adjacent but radically different forest types have very
different canopy faunas but inadequate sampling does not
allow any even remotely accurate estimation as yet of the
extent of ‘turnover’ in moving from one forest type to
another, or whether this is higher or lower than species
turnover in the forest’s lower strata.
29
Species Inventory
In sum, quite enough is known to indicate that high local
Species richness (although not of all groups) and
considerable patchiness at quite a small scale are typical of
tropical forest canopy arthropod communities. How large a
contribution canopy-dwelling species or species that are
present in canopy samples (not exactly the same thing)
make to overall arthropod species richness at one site is less
clear. The contribution made by canopy species to faunas at
regional and other scales is even less well understood,
despite claims that the canopy is where maximum tropical
biodiversity occurs (Erwin, 1990).
Against this background, it is rather surprising that
speculations as to the number of species of arthropods that
might be found overall in the canopies of tropical forests
(Erwin, 1982, etc.) have come to occupy centre stage in
recent general discussion (May, 1988, 1990; Stork, 1988;
etc.) of the possible magnitude of the global species
inventory. At the same time, and stemming from the view
that tropical forest canopies harbour an unparalleled
diversity of life, suggestions that the global species total for
terrestrial arthropods alone may be as high as 50 or even
100 million have also been widely reported, and have found
expression in a number of reports concerned with the
conservation of biotic diversity (Wolf, 1987; Reid and
Miller, 1989; National Science Board, 1989; etc.). The
attention paid to these suggestions perhaps justifies a closer
look at data that may give some hints as to the likely
richness of tropical forest canopy arthropod assemblages.
Tropical forest canopies: reassessment of the evidence
Critical examination of the available data (many of them
still unpublished) might usefully begin with some evaluation
of how fully the richness of canopy arthropod assemblages
is reflected in samples that are routinely studied. Most of
the significant data points come from insecticide fogging
studies. The proportion of species that might be expected to
be obtained by this technique has been the subject of some
discussion (Adis et al., 1984; Erwin, 1990; Stork, 1991;
etc.), but without firm conclusions being reached. However,
restricting attention to adult stages only, we know that some
species that mine or burrow within living or dead plant or
fungal tissue and some of the fauna of suspended litter and
soil are poorly collected by fogging, as are certain
arthropods that are firmly attached (e.g. scale-insects) to
leaf surfaces, along with an uncertain proportion of the
larger species of some groups that may escape capture by
flight. On the other hand, species that are present as
‘tourists’, most of them presumably resting on exposed
surfaces or in flight, seem to be well sampled locally.
Characteristically, their pattern of occurrence in the canopy
is patchy and unpredictable, with the result that tourist
species accumulate steadily as sample size is increased. A
good number of groups (e.g. ladybirds, ants, adult psyllid
bugs, etc.) seem to be sufficiently well sampled by fogging
that results give an accurate impression of the relative and
even absolute abundance of individual species, as well as a
good account of which species are present.
However, canopy samples obtained by means other than the
application of insecticides reveal that a proportion of true
canopy species are not or are not readily taken by fogging.
The most telling evidence for this comes from studies
1. Biological Diversity
(Hammond, 1990; Hammond and Stork, unpublished)
where canopy fogging has been carried out in tandem with
additional extensive sampling of both canopy and lower
forest strata by other means. In such instances we find a
certain number of species well represented in, for example,
baited traps or interception traps placed in the canopy, but
absent from traps of the same type operated at ground level
as well as from fogging samples.
Ignoring the proportion of species (probably rather small)
that are not well sampled by the technique, how much
fogging is necessary to give a reliable picture of the size of
a local canopy arthropod community, and how are its
components distributed? A number of studies in both
temperate and tropical countries suggest that, with an
appropriate pattern of sampling (including adequate seasonal
coverage) relatively few trees or quadrats may be needed.
Particularly good evidence on this point is emerging from
the results of a fogging programme carried out in a
relatively uniform tract of lowland tropical forest in
Sulawesi (Hammond and Stork, unpublished). In this study
a number of samples, covering all seasons, were taken from
each of 20 different 12 x 12m quadrats distributed through
a 500ha study area. A strong indication that a representative
sample of the canopy insects present in the study area was
obtained is furnished by the rate at which species
accumulated with sampling effort (see Fig. 4.3).
How near are we to determining the proportion of all
arthropod species present in a given tropical forest that are
likely to be taken by canopy fogging, and is this more or
less a constant? If canopy samples are to be used as a
means of directly estimating overall species richness of a
forest, either locally or at a larger scale, it is clearly vital
that the relationship between numbers of species present in
the canopy and the number of species found overall be
roughly understood. If canopy samples are to be used for
comparing local species richness directly it would obviously
be helpful if proportions varied little from one place to
another. Finally, if global figures for arthropod species
richness are to be derived from canopy fogging data (see
below) these will be on a particularly sure basis if the
number of species present in canopy samples is a very high
as well as constant and a known proportion of the whole.
That this is the case, for neotropical forests at least, has
been asserted by Erwin (1991) who in earlier work (1982)
suggested that canopy arthropod communities were at least
twice as rich overall as those of the forest strata below.
Working from first principles, this sort of relationship
might seem unlikely. Most of the production of living tissue
in a forest starts off in the canopy, but most of this - fallen
leaves, fruit and wood, insect, bird and other excrement,
and whole fallen trees - ends up forming a rich mosaic of
resources on the forest floor. Not surprisingly, the
abundance and biomass of arthropods is greatly skewed in
favour of the lowest levels in a forest. Strictly comparable
figures for both canopy and forest floor are not available,
deriving as they do from fogging samples for the canopy
(undersampling internal and concealed feeders, etc.) and a
range of different ‘standing crop’ methods for the forest
floor. For example, in neotropical forests investigated by
Adis and Schubart (1985), disregarding the Collembola and
mites which made up 60-80% of the individuals in soil/litter
samples, an average of around 30 times as many arthropods
30
were found, per m”, in the soil/litter layer as in the canopy.
Methods used in studies such as this are known to
undersample small arthropods, mites and springtails in
particular, because of poor extraction from soil and other
substrates, and also ignore or underplay the large
contribution made by significant but patchily distributed
resources such as carrion, fallen fruit, large fungus fruiting
bodies and decaying wood.
Both baited traps and those not involving attractants (e.g.
Malaise traps and window traps) collect far fewer
individuals and species at canopy level than on the ground.
This is a common finding of studies in several countries.
Some tropical studies (e.g. Hammond, 1990), for example,
show a relationship of around three species of Coleoptera
in ground-level Malaise trap samples to one for the same
trapping effort in the canopy. A much higher ground to
canopy ratio is characteristic for some other groups (e.g.
Hymenoptera) and higher ratios all round are generally
found in catches from interception or other traps that do not
favour plant-climbing species.
Apart from temperate forests where the overall proportion
of species present at a site that can be found in the canopy
probably rarely exceeds 20%, the most compelling evidence
for much lower local species richness in the canopy than at
other levels comes from the Sulawesi study already
mentioned (Hammond, 1990), where as complete an
inventory as possible was made of the Coleoptera and some
other insect groups found in the 500ha study area. The
extensive canopy fogging that formed part of the sampling
and inventorying programme produced around 30% of the
beetle species found in total, and around 20% of those
conservatively estimated actually to occur in the study area.
More than three-quarters of the species taken by fogging in
the Sulawesi study were also present in samples of various
types taken at ground level. Analysis of their pattern of
occurrence in all ground and canopy-level samples suggests
that many of these were present in the canopy only as
‘tourists’, and that overall less than two-thirds of species
found in the canopy belong to the canopy fauna proper,
either as ‘specialists’ (species largely restricted to the
canopy) or ‘generalists’ (species found regularly both in the
canopy and at lower levels). Making allowance for canopy
species not obtained by fogging, canopy species proper
amount to at most 20% of the area’s species, of which no
more than half (i.e. probably less than 10% of the total
fauna) may be regarded as canopy specialists.
Results from other palaeotropical and from neotropical sites
suggest that although canopy insect species richness in
tropical moist forests is somewhat variable, it is not
exceptionally low at the Sulawesi site. Somewhat higher
levels of local species richness might be expected, however,
in canopies that contain more tree species and forests in
which canopy, understorey and ground layers are more
clearly demarcated. Data available for temperate forests
suggests relatively weak stratification, a very small canopy
specialist component and a ‘typical’ overall canopy to
ground arthropod species ratio of around 1:10 or more.
Variation is to be expected in tropical forests, with the
lowest ground to canopy ratios most likely to be found
where the ground component is relatively small (e.g. dry
Figure 4.3
so
40
Canopy beetles of various
Species Inventory
Accumulation of beetle species in canopy samples
guilds including tourists
Accumulative Totals of Species
30 |
L
20 : ,
[ Al One canopy herbivore guild
are
10 };—
a} 1 4 4 _L 4 4 4 =: i 1 rt 4 4 4 iL
March Ss July 10 December 15
oO | al | | | ! | | | |
March = July 10 December 15
Number of Samples
Notes: Beetle species in canopy fogging samples from a single tropical site and how these accumulate with increased sampling effort. The upper
curve is for a ‘representative’ selection of 51 species (out of 900 beetle species in the total sample) comprised of 23 ‘regular’ canopy species and
28 that are present in the canopy as ‘tourists’. It shows a steady decrease in increments with sampling effort but no distinct flattening. The inset curve
is for some of the species - the 10 members of a herbivore guild (broad-nosed leaf-chewing weevils that are all either canopy specialists or generalist
species regularly feeding in the canopy) - included in the upper curve. This shows how, with a dataset restricted to canopy species proper, the species
accumulate much more rapidly, in this case reaching a plateau after 5 (out of 15) samples had been taken.
forests) or, conversely, where the canopy component is
high as a consequence of great stratification, as may be the
case in some of the tallest closed-canopy moist forests. The
ratios found for Coleoptera in Sulawesi (about one in five
species belong to the canopy fauna proper, about one in ten
species are canopy specialists) may not be modal for
tropical forests, but further results are needed before any
firm view on what ‘typical’ ratios are can be taken.
Despite the large numbers of arthropod and other species to
be found in tropical forest canopies, there are few data
providing any clear support for the view that the upper
levels of tropical forests are truly the "heart of biotic
diversity". If anywhere, it would seem more likely that this
is to be found on and under the forest floor.
SAMPLING THE HYPER-DIVERSE BUT POORLY
KNOWN
Knowledge of large organisms and some temperate regions
provide an inadequate basis on which to extrapolate with
any confidence to groups and areas that are poorly-known.
Well-established species richness patterns exhibited by
groups such as birds are, of course, a useful starting point
in attempts to gauge better the species richness of less
well-known groups, but there is every reason to suppose
that they provide no more than general guidance.
Well-known organisms are a biased sample of the biota.
Apart from being mostly large, they may also be
31
unrepresentative in many other ways. Vascular plants, for
example, may be much less dependent on surface moisture
levels than many small animals. Butterflies, unlike the
majority of insects, are all essentially herbivorous.
Taxonomic groups, functional groups and ecosystems that
might be expected to make the largest contributions to
global species richness have been briefly surveyed above.
For some of these, there are strong indications of
considerable diversity that is as yet unaccounted for by the
taxonomic inventory, while for others the hints are more
vague. Many more data for these poorly-known groups and
areas of the world are needed for the magnitude of their
contributions to biodiversity to be even roughly assessed.
How some of these data might be gathered and applied to
species richness estimates is discussed below.
What, where and how
Almost any new data on species richness patterns in the
groups discussed in the previous section are likely to prove
useful, but the pace at which our understanding of these
patterns improves will depend heavily on which data we
choose to gather first, and on how economical and effective
the methods are that we adopt.
The questions of what and where to sample and how best to
gather sample data to improve our knowledge of major
species richness patterns has been well reviewed in a recent
1. Biological Diversity
report (Solbrig, 1991) where the need to focus efforts on
high diversity groups and ecosystems is highlighted. More
precise proposals with regard to the choice of sites for
intensive study and the choice of indicator or focal groups
(see below) have been advanced by di Castri et al. (in
press). Clearly, there is an urgent need for better data on all
of the hyper-diverse groups: insects, nematodes, fungi,
bacteria, etc.. However, it is equally clear that we cannot
expect progress to be made at an even rate on all fronts.
The point of departure varies from group to group, as does
the ease and reliability with which good sample data may be
obtained. Some groups are distinctly more tractable than
others, in the sense that large samples may be rapidly and
reliably sorted to species.
To make the most of the considerable effort involved in
gathering species richness data for groups of any size, two
complementary approaches are necessary. The intensive
approach entails in-depth studies, inevitably feasible for
large groups at only a few sites, aimed at establishing the
number of species present as precisely as possible. If
coupled with appropriate quantitative sampling, the process
of intensively inventorying a single site may be exploited to
identify and calibrate methods that are needed for studies of
a more extensive type. Thus, complete site inventories are
needed to furnish the ‘knowns’ against which sampling
methods can be calibrated and more extensive sample data
compared. The actual methods used for inventorying will,
of course, vary from group to group, habitat to habitat, and
biome to biome.
The current emphasis on terrestrial arthropods in
biodiversity research is perhaps to be explained as much by
the general amenability of these animals to study as by the
likely size of their contribution to the global species
inventory.
In extensive studies of hyper-diverse groups it may often
prove necessary to deal with just part of the group rather
than treat it in its entirety. In such instances the ‘indicator’
group or groups chosen need to be as ‘representative’ as
possible. It is also helpful if, in species terms, they
constitute a more or less unvarying proportion of the group
as a whole.
Where to look first if we aim to advance rapidly our
knowledge of species richness patterns in the ultra-diverse
groups is fairly clear. In the marine realm there is an
evident need for many more data from the ocean depths.
For terrestrial organisms in general the most urgent
requirement is for more data from the moist tropics.
Despite their undoubted richness, tropical forests remain the
least well studied of major terrestrial ecosystems.
Kinds of extrapolation
Extrapolation of one sort or another is likely to be
employed at every stage in the process of assembling and
interpreting species richness data on poorly studied groups
of organisms or regions. Although all extrapolative
procedures involve the same assumption: that a ratio
obtaining in a known situation holds in an unknown one,
some kinds of extrapolation may, in practice, be seen to be
more trustworthy than others.
32
For the purposes of this discussion, perhaps the most
important distinction to make is between ratios that are
extrapolated from one site to another and those that are used
to extrapolate across spatial scales. Some of the different
kinds of ratio that may be extrapolated from site to site
have already been mentioned above while discussing the
intensive/extensive approach to obtaining species richness
data. Most commonly, when dealing with sites of a
generally similar type, ratios used will be those relating less
complete (sample/focal group) data to more complete
(inventory/larger group) data. Here, the reliability of
extrapolation will depend in part on how extensively the
ratio has been calibrated, but also of relevance is the notion
of comparing like with like. For example, a ratio that has
been shown to obtain at a series of sites in the moist tropics
might well be considered unlikely to hold at temperate sites.
It goes almost without saying that species richness data for
poorly-known groups that we may wish to use as the basis
for extrapolation will generally relate to single sites, as few
reliable data for larger areas are available. If we start with
single site data and wish to extrapolate to species richness
of such groups at the regional or global level, we face a
dilemma, as the ratios needed can only come from the few
very well-known groups of organisms in which species
number relationships across spatial scales are more or less
established. Such ratios, derived as they are from groups
which in the main may be expected to have quite different
species turnover rates, should be used only with the greatest
caution.
It is, of course, possible to extrapolate directly from species
richness data for a single site or even a single sample to
species richness at the ecosystem, regional or global level.
Naturally enough, approaches that offer the possibility of
moving from sample or site figures to global figures in a
single step are tempting to use. However, given its
inevitably speculative nature, extrapolation in this way is
probably best avoided. The limitations of methods that
involve empirical species richness relationships between
very different groups of organisms (e.g. vascular plants and
insects, butterflies and nematodes), host specificity levels,
and proportions of species remaining undescribed have
already been discussed. In some instances, ratios made use
of (e.g. host: parasite species numbers) are likely to be
extremely poorly calibrated. In most cases, the extrapolation
from site to region or globe involves the essentially unsafe
(and often unstated) assumption that the relationships used
scale evenly (see May, 1990).
NEW DATA ON TROPICAL INSECTS AND WHAT
THEY CONVEY
It is widely assumed that insect species outnumber all
others. The belief is not without some foundation, as more
than half of all described species are insects, and it is
evident that at least several times as many remain
undescribed. Ultimately, however, the question of the size
of the contribution that insects make to the global species
inventory is not to be settled by data on the insects
themselves. A much improved understanding of
microorganismal diversity and a better appreciation of
species richness in groups such as the fungi and nematodes
is needed for the insect contribution to be seen in
perspective. This said, the insect part of the equation is a
matter of obvious interest, particularly if we concede that an
approximate answer to the question of how many insect
species there are is within reach.
In comparison with other speciose groups such as
nematodes or mites, knowledge of tropical insects is
relatively advanced. Although the actual evidence remains
fragmentary and anecdotal in the main, it has long been
recognised that the tropics, and moist tropical forests in
particular, contain far greater numbers of species than
extra-tropical regions. Arguably, therefore, a reasonably
accurate estimate of the number of tropical insect species
would provide a good indication of the scale of insect
species richness overall. For some of the smaller and
best-known insect groups, such as butterflies and
dragonflies, tropical species richness patterns are, in fact,
rather well understood. The same cannot be said of the
largest insect groups, although enough is known concerning
a range of family-level taxa to suggest that the proportional
representation of these groups (Coleoptera, Diptera and
Hymenoptera) in the tropics may differ significantly from
that in well-studied parts of the temperate regions.
New quantitative data, including a number not yet referred
to in print, are beginning to both broaden and give greater
precision to our understanding of tropical insect species
richness and how it is distributed. However, few hard data
on the number of species of any of the major insect groups
to be found at individual tropical sites have yet emerged.
Only for the very best-known groups, such as butterflies, is
there any sound appreciation of turnover rates and the
relationship between single site and regional species
richness.
In spite of these difficulties, two datasets concerning the
number of species of major insect groups present in large
samples taken at moist tropical sites have already been used
(Erwin, 1982; Hodkinson and Casson, 1991) to generate
estimates for total tropical and also global insect (or
arthropod) species richness. The estimates produced from
these now widely quoted studies, both of them involving
explicit assumptions, but with regard to ratios of very
different kinds, are strikingly divergent, with Hodkinson
and Casson arriving at a figure of around two million for
insects globally and Erwin at a figure of 30 million for
arthropods in the tropics alone. If correct, the first figure
implies that around half of all insect species have already
been described, while the second would suggest that
undescribed insect species outnumber those described by a
factor of 30 or more. However, not too much significance
need be read into the discrepancy between the results, as
both approaches entail the use of ratios that are essentially
uncalibrated. Recognising this, Erwin’s (1982) original
calculations have been tentatively reworked by others (e.g.
Stork, 1988; May, 1990), illustrating well how ostensibly
reasonable but different assumptions will produce widely
varying results from the same chain of reasoning. The same
applies, if with less force, to Hodkinson and Casson’s
calculations (see below).
Hodkinson and Casson use a single data point - the number
of species of bugs (Hemiptera sensu Jato) in samples from
the Dumoga area of N. Sulawesi, Indonesia. They suggest
418)
Species Inventory
that the bug samples studied "contained a significantly high
proportion of the species present", but there is good reason
to suppose that the recorded total of 1,690 species
represents a considerable underestimate. However, for the
first of the two separate calculations employed by
Hodkinson and Casson, the extent to which their data
accurately reflect the size and composition of the bug fauna
of their study area is not directly relevant. They begin by
estimating the ratio of undescribed to described species in
the Dumoga sample of bugs and then, treating this as a
subsample of the world bug fauna, extrapolate directly to a
global figure for the group. Only two considerations are of
significance here: the accuracy of the undescribed to
described ratio for Dumoga bugs, and whether the Dumoga
sample is in fact representative in global terms. On the
second count, we lack the data to make any reasonable
judgement, but with regard to the first it is clear that the
estimates on which the ratio is based, as might be expected,
are in no way precise. In fact, the figure of 62.5 for the
percentage of species undescribed could well turn out to be
rather conservative.
The second line of attack adopted by Hodkinson and Casson
begins with the number of undescribed species of Hemiptera
(see discussion above) considered to occur in the Dumoga
area (i.e. 62.5% of 1,690 = 1,056) and the ostensibly
empirical relationship between this and the number of tree
species found there, estimated to be around 500. Direct
extrapolation to the tropics as a whole (with an estimated
50,000 tree species) yields a figure of 105,600 undescribed
tropical bug species. Added to the 81,700 species of bugs
already described, this furnishes a total of 187,300, no
allowance being made for undescribed extratropical species.
It should be noted that the relationship presumed to exist
between the numbers of bug species and numbers of tree
species present in a given area includes the hidden
assumption that this scales evenly, that is to say that an area
containing, for example, 5,000 tree species may be
expected to contain 10 times as many (rather than 5 or 20
times as many) bug species as an area with 500 tree
species. This problem of scaling is as relevant to empirical
relationships of the type considered here as it is to those
based on host-specificity (see discussion in May, 1990).
For both sets of calculations Hodkinson and Casson scale
up to global insect species overall by using figures of 7.5%
or 10% for the proportion of the world’s insects that are
Hemiptera. The first of these figures represents the
proportion of described insects that are Hemiptera, more
reasonably put at around 8.5%, and the second is the
proportion of insect species in Bornean canopy fogging
samples that are bugs. Both are probably over-estimates.
Bugs, like several other mainly plant-associated groups, are
known to be over-represented in fogging samples; for a
number of reasons, including their taxonomic apparency, it
may be reasonable to assume that bugs are proportionately
better described than the insects as a whole. Taking a figure
of 5% (rather than 7.5% or 10%) as the proportion of
insects that are bugs and applying this to the revised bug
estimates produced above, we see that it is possible to reach
figures for world insects in the range 6.5 to 11 million
rather than the two million or so that Hodkinson and
Casson conclude with.
1. Biological Diversity
The ostensible basis for the estimate of 30 million tropical
arthropods obtained by Erwin (1982) is an interesting study
of the beetles (of some 1,200 species) obtained by fogging
the canopies of 19 individual trees of the neotropical species
Luhea seemannii (Erwin and Scott, 1980). In the light of
how little is known of insect species:tree species
relationships, this might seem an unlikely source for an
estimate of tropical arthropod species richness. However,
closer examination of the chain of reasoning adopted by
Erwin reveals that the data obtained from the field on Luhea
insects play a relatively minor part in the calculations. Of
much greater significance in terms of the results are two
major assumptions that are unrelated to the field data. The
first of these, and one which we are far from being in a
position to test concerns average levels of host-specificity in
tree-dwelling tropical insects (see also May, 1990). The
second assumption, one that, at least at the local level, is
much easier to test, concerns the proportion of tropical
forest species that are to be found in the canopy. Other
factors involved in Erwin’s chain of argument, including the
proportion of canopy arthropods that are beetles, and the
number of species of tropical trees, are less problematic, as
the figures used may reasonably be expected to be of the
right general order. It should be added that further implicit
rather than explicit assumptions that relate to problems of
scaling (see discussion in May, 1990) are involved.
The role played by the estimate of 163 for the number of
beetle species specialised on the average species of tropical
tree in Erwin’s estimate is crucial. Essentially, it is this that
generates the very high figure for tropical insect species
richness that eventually emerges from his chain of
calculations. Unfortunately, although there are good reasons
to suppose that the degree of host-specificity exhibited by
tropical canopy insects is generally low, there are few data
that give even a hint as to what actual levels of
host-specificity might be. More importantly, and as has
already been noted, the use of host-specificity data for
species richness calculations is beset with problems (see
discussion in May, 1990). Even in the British Isles, where
the host ranges and preferences of canopy-dwelling insects
are relatively well documented, specificity data are far too
imprecise to be used for any calculation of the number of
tree-associated insect species.
Bearing these limitations in mind, reworking of Erwin’s
calculations may be viewed as of little practical value.
However, it should be noted that truly staggering numbers
are generated if the ratio of tropical canopy beetle species
to tropical beetle species overall is revised in the light of
findings discussed above. If the 1:4 or so canopy to total
ratio found to obtain in Sulawesi is substituted for Erwin’s
2:3, but all else in Erwin’s chain of calculations is left as it
is, we arrive at an estimate for tropical forest arthropods
alone of around 100 million, rather than 30 million. If we
should conclude, reasonably enough in view of what is
known of tropical canopy insects, that beetles are typically
less than 40% of canopy arthropod species, let us say 25%
(see Stork, 1987), the estimate for tropical arthropods rises
again to approaching 200 million.
Some of the relationships used by Erwin are important ones
for almost any kind of estimates of global insect species
richness that we might envisage, and some of these, for
34
example the proportion of tropical forest beetles that are'to
be found in the canopy, are also amenable to test. However,
this is far from true for the key relationship that Erwin
employs, concerning numbers of beetle species that are
effectively specialised on individual species of tree. In fact,
it would seem likely that only when we know most of the
answers that we are actually seeking, i.e. the number of
species of insects to be found in the tropics and how many
of them are found in the canopy, will we be in a position to
start gaining some idea of how many are exclusively
associated with the average tropical tree species.
The methods of estimating tropical insect species richness
used by Erwin on the one hand and Hodkinson and Casson
on the other have been discussed in some detail here with
the intention of stressing the problems involved in the
short-cut approach. Any extrapolatory route, from sample
or inventory data to a summary for the tropics as a whole,
that avoids the explicit use of ratios concerning relative
species richness at different spatial scales is bound to be
tempting. However, if the alternative is to invoke
relationships that cannot be calibrated, the temptation is
perhaps best avoided.
The valuable datasets (Casson, 1988; Erwin and Scott,
1980) on which the Hodkinson and Casson and Erwin
estimates discussed above were based are just a part of a
whole crop of new data that have recently become available
for tropical insects. Although most results pertain to rather
narrow taxonomic groupings, they are nevertheless leading
to a steady improvement in our overall understanding of
such questions as altitudinal gradients in species richness,
species turnover at small spatial scales, and the contribution
made by elevational assemblages and pronouncedly different
but adjacent forest types to species richness at the level of
the ‘extended site’.
Data of a particularly extensive type have come from one
recent large study based on an area of moist tropical forest
in northern Sulawesi, Indonesia. The work of analysing
results is still in progress, but many data concerning local
species richness of beetles (Hammond, 1990) have already
become available. The full dataset for beetles includes the
results of quantitative sampling by a variety of means
through all seasons of one year, as well as an inventory of
species found within the principal study area (SO0ha of
relatively uniform lowland forest). Valuable if less
comprehensive data for several other insect groups, e.g.
Hemiptera (Casson, 1988) and Hymenoptera (Noyes, 1989)
are also available. The data from this study offer the
possibility, for the first time, of (1) establishing a figure for
overall local species richness of some major insect groups
at a tropical moist forest site, (2) assessing what proportion
of species is found in the canopy as opposed to lower layers
(see above), and (3) of calibrating a range of sampling
methods against knowns (total inventory results) in a
tropical forest setting. Finally, the detailed sample data and
inventory provide a comprehensive enough picture of the
assemblage of insects present that, with sufficient general
knowledge of their biology, it is possible to assess the
proportions that belong to different functional groups, and
that are associated with particular microhabitats and the
various forest strata. The biases of various sampling
methods with respect to these and other characteristics, such
as body size and taxonomic group membership, may also be
determined.
The findings of most direct relevance to overall tropical
insect species richness to emerge so far from this study are:
© Species richness of Coleoptera at this tropical site, at
scales of lha up to around 500ha is some five times
greater than the average for a range of temperate forest
sites. The species richness of Hemiptera, in relation to
temperate sites, may be of the same general order, while
that of Lepidoptera and Hymenoptera is also higher than
in temperate forests, but by a less certain factor
(probably between two and four)
e The numbers of species of some major insect groups and
of insects overall that are found in the canopy are low
compared with numbers found at ground level
e For an equivalent intensity and pattern of sampling, some
of the sampling methods used obtain the same proportion .
of species present as they do at comparable sites in
temperate regions (see Fig. 4.4).
In the long term, the last of these findings may turn out to
be the most significant. Following calibration against the
Sulawesi site inventory, simple ‘sampling packages’ that
have already been shown reliably to reflect local species
richness of Coleoptera and/or other major insect groups at
‘known’ temperate sites, might reasonably be expected to
provide a good indication of species richness at other moist
tropical sites. In fact, a number of trials of these sampling
packages at a range of sites in the Indo-Australian and New
World tropics have now been made. Assuming that the
results being obtained (Hammond, unpublished) are reliable,
they suggest ratios for the number of Coleoptera species
between the tropical sites investigated and average
temperate forests, that vary, except for one small tropical
island with substantially lower beetle species richness, from
around 3:1 to 8:1.
New data on the overall species richness of major groups at
single well-defined sites make an obvious contribution to
our general understanding of the pattern of insect species
richness in the tropics. Furthermore, if accurate, they
provide us with the essential base-line from which improved
estimates of tropical insect species richness might eventually
grow. For the moment, our poor understanding of species
turnover in the tropics means that we have little to go on,
if we wish to use single site data for extrapolation to
regional or global figures.
Of course, starting with the ratio of five beetle species at a
moist tropical site to one at a temperate site, crude
extrapolation to a global insect species total is possible, but
to do this it is necessary to make a series of major
assumptions, not the least of which concern the proportion
of insect species that are beetles and, as we have noted,
species turnover rates. For a start, we may repeat the
simple extrapolation made earlier on, based on the
assumption that we are already able roughly to estimate the
number of extratropical beetle species. If we take 400,000
as the number of described beetle species, and assume (no
good count is available) that roughly 50% of described
beetle species are from extratropical regions, and make an
educated guess that around 50% of extratropical species
35
Species Inventory
have been described, an overall ratio of five tropical beetle
Figure 4.4 Beetle species richness:
tropical vs temperate
Trepical Forest
400 + ae
Temperate Forest
Accumulative Totals of Species
Notes: Comparison of beetle species richness in comparable sets of
samples from single tropical and temperate sites, showing a
relationship of around 5 to 1. The graph depicts accumulative numbers
of species over time collected by representative single Malaise traps of
modest size (see Hammond, 1990). Tropical data are for moist lowland
forest in N Sulawesi and temperate data for mixed deciduous woodland
in southern Britain (Hammond, unpublished). Traps chosen for
illustration are those producing total beetle species nearest to the means
of 412 per trap for Sulawesi (9 traps) and 83 for Britain (5 traps).
species to one extratropical species yields a world total for
beetles of 2.4 million, of which two million are tropical and
0.4 million extratropical species. If we then take the
proportion of insect species that are beetles (see below) to
be 33%, the figure we reach for insect species globally is
around 7.2 million. This, of course, involves the dubious
assumption that the tropical to temperate ratio scales evenly
from site upwards, in both tropical and extratropical
regions. Assuming much higher species turnover rates in
the tropics, but bearing in mind that the extratropical
component includes contributions from broad latitudinal
bands in both southern and northern hemispheres, a tropical
turnover ‘factor’ may be brought into play. If we take this
to be (say) 1.3 and apply it to the calculation already made
our figure for insect species worldwide is 9.4 million.
An alternative approach is to take the beetle species total
for the Sulawesi site, and scale up directly to a figure for
the tropics as a whole, using available data on tropical
species turnover for relatively well-known groups as a
rough guide. Using information patched together from many
groups, including the best-known families of beetles
themselves, an extrapolation may be made from the
Sulawesi site inventory of 6,000 or so beetle species to
28,000 for the northern part of Sulawesi, to 70,000
(Sulawesi as a whole), 700,000 (Asian tropics) and finally
1.8 million beetle species for the entire moist tropics. Using
the same figure for extratropical beetles as before, we reach
a global beetle species figure of 2.3 million beetle species
and, assuming (as before) that beetles comprise 33 % of the
global insect species inventory, 6.9 million insect species
worldwide.
Finally, we might compare these results with those obtained
by a Hodkinson and Casson type approach to the Sulawesi
beetle data. In fact, no estimate is available for the
proportion of species undescribed in the sample as a whole,
1. Biological Diversity
Figure 4.5 Major groups of organisms:
total
Vertebrates (2.7%)
Nematodes (0.9%)
Molluscs (4.2%)
Other invertebrates (4.0%)
Other arthropods (1.2%)
Crustaceans (2.4%)
Arachnids (4.5%)
Other insects (8.9%)
Diptera (7.1%)
Hymenoptera (7.7%)
described species as proportions of the global
Plants (Embryophytes) (14.3%)
Algae (2.4%)
Protozoans (2.4%)
Fungi (4.2%)
Bacteria (0.2%)
Viruses (0.3%)
Coleoptera (23.8%)
: =
ARTHROPODS
Lepidoptera (8.9%)
Notes: Proportions of major groups of organisms in terms of described species (estimated to total approximately 1.7 million). Groups included in
the pie-chart are those considered likely to contain in excess of 100,000 species when as yet undescribed species are taken into account, along with
vertebrates for comparison. Numbers of described species used in this diagram are those given in Table 4.3, with the exception of plants for which
an earlier lower estimate of 240,000 was used.
but assuming this (on the basis of a small and probably
unrepresentative sample) to be 75%, we generate a world
figure of 1.6 million species for Coleoptera and, using the
33% formula from above, one of approaching five million
for insect species worldwide.
Of course, all of the more significant ratios used in these
simple calculations derive, at best, from informed guesses,
but they are not simply plucked from the air. First-hand
experience of how heterogeneity manifests itself at very
small scales at tropical sites, and a feel for the extent of the
contribution made by the different elements (e.g. elevational
assemblages and different forest types) involved at more
‘extended’ sites may provide particularly useful guidance.
Knowledge of vicariance patterns, especially as they differ
between the three major tropical regions, may also be of
considerable assistance. Finally, an awareness of the biases
of various sampling methods, and the many factors that
influence how well sampled and studied particular groups
are likely to be, will be of great help when attempting to
grasp the significance of fragmentary data.
The more important ratios used, those concerning the
proportional representation of the major insect groups in
terms of species and tropical to extratropical relationships,
in the simple extrapolations made above were derived by
patching together small fragments of data from many
36
sources. The conclusions reached and assumptions used in
reaching them cannot be detailed here, but it should be
mentioned that higher tropical to temperate ratios were
assumed for Coleoptera, Hemiptera and Lepidoptera, as
opposed to Diptera and Hymenoptera (see Gaston, 1991a).
The relative species richness of what seem certain to be the
three largest insect groups was based on separate
assessments of their possible overall species richnesses in
both tropical and extratropical regions. ‘Working figures’
arrived at for the percentage of insects overall that are
Coleoptera, Hymenoptera and Diptera in extratropical
regions were 25%, 30% and 30% respectively, while those
for the tropics were 35%, 27% and 20%, yielding (if we
assume a 5:1 tropical to extratropical ratio for beetle
species) overall working figures of 33% Coleoptera, 27.5%
Hymenoptera and around 22% Diptera.
PROSPECTS FOR IMPROVED SPECIES RICHNESS
ESTIMATES
Currently available estimates of species richness for all but
the best-known groups such as birds, and best-known
regions such as northern Europe, all involve substantial
margins of error. By simple extrapolation from the
well-known, only a very rough idea may be gained of how
many species exist overall. The many uncertainties,
especially with respect to microorganisms, make an upper
Species Inventory
Figure 4.6 Major groups of organisms: possibly-existing species as proportions of the
global total
Vertebrates (0.4%) plants
Nematodes (4.0%)
Molluscs (1.6%)
Other Invertebrates (1.1%)
Other arthropods (0.5%)
Crustaceans (1.2%)
Arachnids (6.0%)
Other Insects (4.0%)
Diptera (12.9%)
ARTHROPODS
Hymenoptera (19.3%)
(Embryophytes) (2.4%)
Igae (1.6%
Protozoans (1.6%)
Fungi (8.0%)
Coleoptera (24.9%)
Lepidoptera (3.2%)
Notes: Possible proportions of major groups of organisms based on conservative estimates (see Table 4.3) providing a total for all groups of
approximately 12.5 million species. All groups considered likely to contain in excess of 100,000 species are picked out in the pie-chart, along with
vertebrates for comparison.
bound to the size of the global species inventory particularly
difficult to establish. Despite numerous indications that this
could be very great, claims that extant species number
many tens of millions or even more can not be supported,
for the moment, by any firm evidence. However, a lower
bound to the global figure is much easier to set, and the
available data, some of them discussed above, suggests that
this might safely be put at a level considerably higher than
the current described species total (approaching two
million), perhaps at around eight million. The ‘working
figure’ adopted here of 12.5 million species for the biota as
a whole (see Fig. 4.6), arrived at by examining the data for
each major group separately, is an avowedly conservative
one.
In a situation where the most species-rich groups are at the
same time the least known, an unwillingness to take into
account anything but incontrovertible evidence is always
likely to result in underestimation, as the record of early
attempts to estimate the scale of global species richness well
illustrates. Nevertheless, if we are to have any confidence
in species richness estimates, there is no real alternative to
working forwards by steadily enlarging the area of knowns.
While new observations concerning little-known taxonomic
groups and poorly explored habitats continually alert us to
additional possibilities of as yet unassessed species richness,
it would be naive to make too much of each and every
anecdote.
37
To speed up the rate at which our understanding of species
Tichness patterns and the overall dimensions of global
biodiversity grows, it will be necessary to identify key
questions and, if feasible, turn our attention first to them.
The most obvious general line of attack is to focus efforts
on the groups of organisms and parts of the globe that seem
most likely to make the greatest overall contribution to the
species inventory. New and pertinent data are needed for all
of the ultra-diverse groups, but quicker and more substantial
returns for efforts made are to be expected from some
groups rather than others. If the main emphasis of this
section has been on terrestrial arthropods, it is not because
these animals (however numerous their species) can supply
all of the answers, but rather because answers to key
questions concerning their patterns of species richness are
seen to be distinctly and not too distantly attainable.
Perhaps the greatest need is for good sample data on
microorganisms and fungi. Because of seasonality and
difficulties in detecting and/or culturing small species,
exhaustive inventories may not be achievable but, in line
with recent recommendations, these should be attempted at
representative sites in the major biomes. If tropical to
temperate species richness ratios are to be established for
these groups, there will be a need to develop sampling
methods and protocols that allow reliable comparison
between sites without a complete inventory being taken.
1. Biological Diversity
For nematodes there is a pressing need for data on the
species richness of free-living forms in both marine and
terrestrial environments, including the moist tropics. Sample
data that allow some estimation of species turnover at least
at relatively small scales (i.e. in the m? to km? range) are a
particular need, while any results concerning the less easily
addressed problem of turnover at larger scales would be of
great value. As in the case of microorganisms, advances in
both the theory and practice of species recognition and
discrimination will be needed if data gathered are to be
truly informative.
In the case of terrestrial arthropods, the more tractable
groups that are also large and ‘representative’ (e.g.
Coleoptera) may be expected to receive considerable
attention. For some of these groups, sampling programmes
at various tropical and other sites, are already well
advanced. While there is a need for the analysis of results
already obtained to be speeded up, this should not be
allowed to stand in the way of the application of the best of
the methods so far developed at many additional sites.
For some of the major terrestrial arthropod groups, e.g.
Diptera and Acari (mites), data on the numbers of species
to be found at any one location in the moist tropics remain
extremely limited and largely anecdotal. Reasonably reliable
estimates of the species richness of these groups at single
tropical sites are eminently attainable, and the acquisition of
the appropriate datasets is a particular priority. Another
clear need is for a better understanding of the proportional
representation, in species terms, of the major terrestrial
arthropod groups at single sites, and how this varies from
region to region.
A separate agenda of research is needed for the
investigation of species richness patterns in the marine
realm. Here, data from the ocean depths remain too
fragmentary for any confident estimation of the contribution
that this ‘new frontier’ might make to marine or overall
global species richness. There is a particularly urgent need
for results that give some idea of species turnover in
deep-ocean sediment assemblages, especially at the larger
spatial scales.
Attention has been directed in this section almost entirely
towards species, which for sound theoretical as well as
operational reasons are often considered "central to the
concept of biodiversity" (Reid and Miller, 1989). However,
it should be stressed that a species count falls far short of
any full assessment of biotic diversity, which expresses
itself at a number of levels, from genes to ecosystems
(Solbrig, 1991).
Our perception of the full dimensions of biotic diversity
remains very hazy, but there is much of an immediate
nature that can be done and is being done to remedy the
situation. Indeed, there is every reason to suppose that
advances in our understanding of some significant species
richness patterns will be made very rapidly. Of course, we
shall not get to know, even approximately, how many other
species we share the planet with overnight, but we may
reasonably expect our global species estimates to be made
38
with steadily increasing confidence and precision.
References
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London
Adis, J., Lubin, Y.D. and Montgomery, G.G. 1984. Arthropods from
the canopy of inundated and terra firma forests near Manaus,
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technique. Studies on Neotropical Fauna and the Environment
19:223-236.
Adis, J. and Schubart, H.O.R. 1985. Ecological research on
arthropods in central Amazonian forest ecosystems with
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Golley, F.B. (Eds), Trends in Ecological Research for the 1980s.
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Arnett, R.H. 1967. Present and future systematics of the Coleoptera in
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Arnett, R.H. 1985. American insects: handbook of the insects of
America north of Mexico. Van Nostrand Reinhold, New York.
Barnes, R.D. 1989. Diversity of organisms: how much do we know?
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Brusca, R.C. and Brusca, GJ. 1990. Invertebrates.
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Casson, D. 1988. Studies on the Hemiptera communities of
Dumoga-Bone National Park, Sulawesi. M.Phil. Thesis. Liverpool
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di Castri, F., Vernhes, J.R. and Younes, T. (in press). A proposal for
an international network on inventorying and monitoring of
biodiversity. Biology International, Special Issue 27.
Colinvaux, P. 1980. Why Big Fierce Animals Are Rare. Pelican Books,
London.
Eliot, J.N. and Kawazoé, A. 1983. Blue Butterflies of the Lycaenopsis
Group. BM(NH) London.
Erwin, T.L. 1982. Tropical forests: their richness in Coleoptera and
other arthropod species. Coleopterists’ Bulletin 36:74-75.
Erwin, T.L. 1990. Canopy arthropod biodiversity: a chronology of
sampling techniques and results. Revista Peruana de Entomologia
32:71-77.
Erwin, T.L. 1991. How many species are there? Revisited.
Conservation Biology 5:1-4.
Erwin, T.L. and Scott, J.C. 1980. Seasonal and size patterns, trophic
structure and richness of Coleoptera in the tropical arboreal
ecosystem: the fauna of the tree Luehea seemannii Triana and
Planch in the Canal Zone in Panama. Coleopterists’ Bulletin
34:305-322.
Frank, J.H. and Curtis, G.A. 1979. Trend lines and the number of
species of Staphylinidae. Coleopterists’ Bulletin 33:133-149.
Gaston, K.J. 1991a. The magnitude of global insect species richness.
Conservation Biology 5:283-296.
Gaston, K.J. 1991b. Body size and probability of description; the
beetle fauna of Britain. Ecological Entomology 16:505-508.
Gaston, K.J. 1992. Estimates of the near-imponderable: a reply to
Erwin. Conservation Biology 5:564-566.
Goodwin. 1986. Crows of the World. BM(NH), London.
Grassle, J.F. 1989. Species diversity in deep-sea communities. TREE
4:12-15.
Grassle, J.F.
41:464-469.
Grassle, J.F., Laserre, P., McIntyre, A.D. and Ray, C.G. 1991.
Marine biodiversity and ecosystem function. Biology International,
Special Issue 23:i-iv, 1-19. TUBS, Paris.
Hammond, P.M. 1990. Insect abundance and diversity in the
Dumoga-Bone National Park, N. Sulawesi, with special reference
to the beetle fauna of lowland rain forest in the Toraut region. In:
Knight, W.J. and Holloway, J.D. (Eds), Insects and the Rain
Forests of South East Asia (Wallacea). Royal Entomological
Society, London. Pp.197-254.
Hawksworth, D.L. 1991. The fungal dimension of biodiversity:
magnitude, significance and conservation. Mycological Research
95:641-655.
Sinauer,
1991. Deep-sea benthic biodiversity. Bioscience
Hodkinson, I.D. and Casson, D. 1991. A lesser predilection for bugs:
Hemiptera (Insecta) diversity in tropical rain forests. Biological
Journal of the Linnean Society of London 43:101-109.
Lawrence, J.F. 1982. Coleoptera. In: Parker, S.P. (Ed.), Synopsis and
Classification of Living Organisms. McGraw-Hill, New York. Pp.
482-553.
May, R.M. 1988. How many species are there on earth? Science
241:1441-1449.
May, R.M. 1990. How many species? Philosophical Transactions of
the Royal Society B330:293-304. ;
Mayr, E., Linsley, E.G. and Usinger, R.L 1953. Method and
principles of systematic zoology. McGraw-Hill, New York.
National Science Board 1989. Loss of Biological Diversity: a global
crisis requiring international solutions. National Science
Foundation, Washington, DC.
Noyes, J.S. 1989. The diversity of Hymenoptera in the tropics with
special reference to Parasitica in Sulawesi. Ecological Entomology
14:197-207.
Poinar, G.O. 1983. The Natural History of Nematodes. Prentice Hall,
Englewood Cliffs, NJ.
Reid, W.V. and Miller, K.R. 1989. Keeping options alive. The
scientific basis for conserving biodiversity. World Resources
Institute, Washington, DC.
Sibley, C.G. and Monroe, B.L. Jr 1990. Distribution and Taxonomy
of Birds of the World. Yale University Press, Yale.
Simon, H.R. 1983. Research and publication trends in systematic
zoology. Ph.D. thesis. The City University, London.
4M)
Species Inventory
Solbrig, O. (Ed.) 1991. From genes to ecosystems: a research agenda
for biodiversity. Report of an IUBS-SCOPE-UNESCO workshop,
Harvard Forest, Petersham, Ma. USA, June 27-July 1, 1991.
TUBS, Cambridge, Mass.
Stork, N.E. 1987. Guild structure of arthropods from Bornean rain
forest trees. Ecological Entomology 12:69-80.
Stork, N.E. 1988. Insect diversity: facts, fiction and speculation.
Biological Journal of the Linnean Society of London 35:321-337.
Stork, N.E. 1991. The composition of the arthropod fauna of Bornean
lowland rain forest trees. Journal of Tropical Ecology 7:161-180.
Southwood, T.R.E. 1978. The components of diversity. In Mound,
L.A. and Waloff, N. (Eds), Diversity of Insect Faunas. Symposia
of the Royal Entomological Society of London. 9. Blackwell
Scientific Publications, Oxford.
Valiela, I. 1984. Marine Ecological Processes. Springer Verlag, New
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Wolf, E.C. 1987. On the brink of extinction: conserving the diversity
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Washington, DC.
Abridged from a document provided by Peter Hammond,
Environmental Quality Programme, The Natural History
Museum (London).
Data presented in Table 4.5 retrieved from Zoological
Record Online by BIOSIS, UK.
1. Biological Diversity
5. SPECIES DIVERSITY: AN INTRODUCTION
A BRIEF HISTORY OF DIVERSITY
Knowledge of the history of diversity through geological
time is based on analysis of the fossil record. Because the
fossil record gives only a very incomplete and highly biased
view of the past history of life on earth, the reconstruction
of that history has been, and continues to be, the subject of
great debate. It is generally accepted that the fossil record
can give a reasonable insight into past diversity in terms of
taxonomic richness, particularly at higher taxonomic levels.
However, it is far more difficult to derive other, more
ecologically based, measures of diversity from it, as these
require the reconstruction of palaeoenvironments, a far
more contentious exercise than palaeotaxonomy.
While detailed patterns of taxonomic richness through the
earth’s history remain debatable, the overall outline is
generally accepted. There are believed to have been
relatively few species in total during the Palaeozoic and
early Mesozoic; since then, that is for the past hundred
million years, diversity has increased markedly. This recent
diversification has passed through one major extinction
event, at the Cretaceous-Tertiary boundary, and probably
two minor events since then (see Chapter 16). Apart from
these, the diversification appears to have continued more or
less unabated, with the world apparently reaching its highest
ever level of species richness during the Pliocene and
Pleistocene, when climatic change and the advent of
organised human activity finally halted the process.
Significantly, however, diversity at higher taxonomic levels
does not conform with this pattern, as evinced by the far
higher number of animal phyla present in the early
Cambrian than today (see below).
The early history of Life - the Precambrian
Recent consensus suggests that cellular life on the planet (in
the form of procaryotes, at least some of which were
probably very similar to living cyanobacteria) originated
sometime between 3,900 and 3,400 million years ago
(Mya). The origin of the earliest eucaryotes has proved
difficult to establish, but it is generally accepted that the
Precambrian microfossils known as ‘acritarchs’, which are
recorded as far back as 1400 Mya, are almost certainly the
cysts of marine algae and the earliest known eucaryotes. If
this analysis is correct, then life on earth consisted only of
procaryotes for at least 2,000 million years, or well over
half its history. There is sufficient morphological variation
in the fossil remains to permit some analysis of changes in
diversity of these presumed early procaryotes in the late
Proterozoic era. Vidal and Knoll (1983) have hypothesised
a gradual increase in diversity from 1400 Mya to 750 Mya,
when there was a peak of around 30 taxa in the fossil
record, followed almost immediately by a sharp drop to
around 10 taxa, possibly owing to a period of glaciation.
After this there is an exponential increase in diversity,
corresponding with the start of the Phanerozoic era.
The early Phanerozoic
For many years it was assumed that metazoans
(multicellular organisms with internal organs) originated in
40
the Cambrian era at the base of the Phanerozoic. This is
now known not to be the case, as a wide range of fossil
metazoans is now known from well before this time,
including recognisable arthropods and _ possibly
echinoderms. Most fossils from this time, however, appear
completely unrelated to extant forms, and consist mainly of
enigmatic frond- and disc-shaped soft-bodied animals: the
so-called Ediacaran fauna.
The lower Cambrian marks a dramatic change from this
early fauna, with the sudden appearance in the fossil record
of a wide range of metazoans, many with calcareous
skeletons. It is generally accepted that this represents a
genuine explosion of diversity which took place over only
a few million years, and is not an artefact of the fossil
record. The lower Cambrian thus represents the most
important period of high-level diversification in the history
of animal life on earth. Very many phyla may have existed
at this time, no more than five of which have origins
traceable to before the Cambrian-Precambrian boundary.
These include every well-skeletalised animal phylum living
today (with the possible exception of the Bryozoa),
indicating that virtually no new animal phyla have appeared
during the many subsequent evolutionary radiations.
Perhaps most significantly, no new animal phyla appeared
with the colonisation of land, some 50-100 million years
after the Cambrian radiation.
The Cambrian appears to have represented not only a peak
of diversification but perhaps also a peak of higher order
taxonomic diversity, as suggested by the presence of many
more animal phyla than the 35 or so now extant.
Changes in diversity of marine animal taxa through the
Phanerozoic
Although the number of phyla has decreased markedly since
the Cambrian, diversity at all lower taxonomic levels has
either increased overall or in a few cases remained more or
less level.
The number of orders (of marine animals) present in the
fossil record climbed steadily through the Cambrian and
Ordovician, levelling off towards the end of the Ordovician
to a figure of between 125 and 140, which has been
maintained throughout the Phanerozoic.
The diversity of families represented in the fossil record
shows a similar pattern of increase through the Cambrian
and Ordovician, levelling off at around 500, a figure which
was maintained until the late Permian mass-extinction (see
Chapter 16). This extinction event resulted in the loss of
around 300 families; subsequent to this, family diversity has
increased to the modern level, with a number of temporary
reversals in the form of the series of extinction events
outlined in Chapter 16.
The trend in number of species in the fossil record is even
more extreme. From the early Cambrian until the mid-
Cretaceous, the number of marine species remained low;
since then, that is in the past 100 million years, it has
probably increased by a factor of 10.
Diversity patterns in terrestrial animals
Colonisation of land by animals has occurred many times;
although the oldest body fossils of terrestrial animals date
from the early Devonian, it is generally accepted that the
primary period of land invasion by animals was the
Silurian.
The overwhelming number of described extant species of
terrestrial animals are insects and arachnids. The fossil
record for both these groups is generally scanty.
Some attempt has been made, however, to chart changes in
insect diversity at the generic level. Insects first appear in
the fossil record in the Carboniferous. The number of
genera then increased through much of the Palaeozoic and
first part of the Mesozoic, interrupted by a sharp drop
coinciding with the late Permian mass extinction, and then
levelling off during the late Triassic. Diversity then doubled
during the Cenozoic or Tertiary, coincident with the
radiation of the angiosperms.
The fossil record of terrestrial vertebrates is much better,
particularly that of tetrapods. The bird record is much less
substantial than that for other groups, probably because
their light skeletons have been less frequently preserved.
Terrestrial vertebrates first appear in the fossil record in the
late Devonian. Diversity remained relatively low during the
Palaeozoic, with around 50 families, and actually declined
overall during the early Mesozoic. From the mid-
Cretaceous the number of families started to increase
rapidly, reaching a Recent peak of around 340. Diversity of
genera follows this overall pattern in a more exaggerated
form. These trends are shown in Fig. 5.1.
Figure 5.1 Fossil diversity: terrestrial
vertebrates
Birds
Mammals
#2 Reptiles
i Amphibians
& Ss
8
Number of orders
8
10
421 408 360
286 248 213
Million years
144 65
Source: Adapted from Signor, P.W. 1990. The geological history of
diversity. Annual Review of Ecology and Systematics 21.
Note: Diversity is here measured in terms of number of taxonomic
orders present.
Diversity patterns in vascular plants
It is generally accepted that vascular terrestrial plants first
41
Species Diversity: An Introduction
arose in the Silurian, although some palaeobotanists argue
for a Late Ordovician origin. Diversity increased during the
Silurian, and then more rapidly during the Devonian, owing
to the first appearance of seed-bearing plants, leading to a
peak of over 40 genera during the late Devonian. Diversity
then declined slightly, but started to increase markedly
during the Carboniferous, with at least 200 species recorded
by the mid Carboniferous. Following this, diversity
increased only slowly until the end of the Permian. There
was a minor decrease in diversity at the end of the Permian,
coinciding with or preceding the mass extinction of animal
species, followed by a rapid rebound to previous levels.
Diversity then continued increasing slowly, reaching around
250 species in the early Cretaceous. Starting at the mid-
Cretaceous, diversity began increasing at an accelerating
pace.
This overall pattern masks important changes with time in
the composition of the flora, most notably in the relative
importance of the three main groups of tracheophytes: the
pteridophytes, gymnosperms and angiosperms. The Silurian
and early Devonian are marked by a radiation of primitive
pteridophytes. During the Carboniferous, more advanced
pteridophytes and gymnosperms developed and underwent
extensive diversification. Following the late Permian
extinction event, pteridophytes were largely replaced
(although ferns remain abundant) by gymnosperms which
became the dominant group until the mid-Cretaceous. The
dramatic increase in plant diversity since then is entirely
due to the radiation of the angiosperms which first appeared
in the lower Cretaceous. These trends are shown in Fig.
522
Figure 5.2 Fossil diversity: terrestrial
plants
Flowering plants
2 Gymnosperms
@ Ferns
ee
Number of species
ELE “
421 408 360
286 248 213
Milion years
Source: Adapted from Signor, P.W. 1990. The geological history of
diversity. Annual Review of Ecology and Systematics 21.
Note: Diversity is here measured in terms of number of species
present.
144 & 2
MEASURING BIOLOGICAL DIVERSITY
A central problem in the maintenance of biological diversity
is an assessment of the relative importance, in terms of
diversity, of different areas, habitats or ecosystems. Only
by understanding this can priorities in conservation efforts
be usefully assigned. However, this importance can be
assessed in different, though related, ways. The first, and
most obvious, makes reference to its ‘intrinsic’ diversity, so
that an area with higher diversity is deemed more important
than one with lower diversity. The second attempts an
1. Biological Diversity
assessment of the contribution any given area makes to
theoverall diversity of a given geographic region, such as a
country, continent or, ultimately, to the world overall.
From this perspective, some areas with lower intrinsic
diversity may be more important than others with higher
diversity. This will be discussed further below; see also
Chapters 2 and 15.
Assessments of diversity pose considerable problems, both
practical and theoretical. In the first instance, the concept
of diversity in an ecological context has to be made clear.
Local biological diversity
Species richness
Biological diversity measures for particular areas, habitats
or ecosystems are often largely reduced to a straightforward
measure of species richness. In its most ideal form this
would consist of a complete catalogue of all species
occurring in the area under consideration. In practice this
is clearly unrealistic outside very small areas which will be
of only limited interest in a global context. Even with small
sites, a complete enumeration of all species will be
impossible to carry out if micro-organisms are included.
Species richness measures will therefore in practice be
based on samples. Such samples could consist of complete
catalogues of all species in a particular, generally
taxonomic, group (e.g. all birds, all ferns) or may consists
of measures of species density (i.e. all the species in a
sample plot of standard area) or of numerical species
richness, defined as the number of species per specified
number of individuals or biomass.
Although straightforward measures of species richness may
convey relatively little ecologically important information,
in practice because they are the most easily derived, they
are perhaps the most useful index for comparisons of
biological diversity on a large scale.
Species abundance
From an ecological viewpoint, simple species richness
indices have limited value. More meaningful measures of
diversity take into account the relative abundance of the
species concerned. In general, the more equally abundant
the species in the area or ecosystem under consideration
are, the more diverse it is considered to be. A number of
models have been developed which derive diversity indices
from measures of species abundance. As_ different
mathematical and biological assumptions are made in these
models, they will often generate different diversity
measures from the same sets of data. Thus there is no one
authoritative index for measuring diversity.
Taxic diversity
Furthermore, weight can also be given to the relative
abundance of species in various categories, for example in
different size classes, at different trophic levels, in different
taxonomic groups, or with different growth forms. Thus a
hypothetical ecosystem which consisted only of several
species of primary producers, such as photosynthesising
plants, would be less diverse than one with the same
number of species but which included herbivores and
predators. Similarly, an ecosystem with representatives
42
from four different phyla would be more diverse than one
with representative of only two.
Based on cladistic analysis, a number of taxonomic
diversity indices have now been developed. Some of these
give higher weight to so-called relict groups, that is
taxonomic groups not closely related to other living groups
and consisting of few species; others favour higher
taxonomic groups with large numbers of species. The most
complex measure so far developed is taxonomic dispersion,
which endeavours to select an even spread of taxa in any
given group.
Comparisons of different areas
Once a measure of diversity has been decided upon, it
should be possible to compare the diversity of different
areas. Such comparisons may not, however, be
straightforward.
Diversity measures for ecological entities such as
communities, habitats and ecosystems make the assumption
that these entities are not site-specific, that is that they
occur in essentially the same form over a wide area or ina
number of different places. In practice, species composition
and species abundance are very rarely constant either in
space or in time; thus the existence of communities or
ecosystems definable by species composition is seriously
questioned by many ecologists. This therefore undermines
the extent to which diversity measures derived from
particular sites can be used as a basis for generalisation.
Nevertheless, these ecological concepts still retain
considerable force, even if they cannot be rigorously
defined, and much discussion of biological diversity is
couched in terms of comparisons between different habitats
and ecosystems.
Species/area relationships
The relative diversity of different sites will often depend on
the scale at which diversity is measured. Thus 1m? of semi-
natural European chalk grassland will contain many more
plant species than 1m? of lowland Amazonian rain forest
whereas for an area of, say, 1km* or more this will be
reversed. This is because as an area is sampled the number
of species recorded increases with the size of the area, but
this rate of increase varies from area to area.
A wide range of observations has demonstrated that, as a
general rule, the number of species recorded in an area
increases with the size of the area, and that this increase
tends to follow a predictable pattern, known as the
Arrhenius relationship, whereby:
logS = c + zlogA
where S = number of species, A = area and c and z are
constants.
The slope of the relationship (z in the equation above)
varies considerably between surveys, although is generally
between 0.15 and 0.40, and some surveys do not fit the
relationship at all. This relationship is shown graphically in
Fig. 5.3.
Figure 5.3 A typical species-area plot
2
n
C
Uv
Q
Q ek
° * *
L
o
ee
—E
3
c
ry
4 *¥
1 2 3 4
Log area (ha)
Note: The data are plotted on logarithmic axes resulting in a straight
line graph, the slope of which (z) indicates the rate at which species
number changes in step with changing area.
The most widely quoted generalisation from this finding is
that a ten-fold reduction in an area (i.e. loss of 90% of
habitat) will result in the loss of from c. 30% (with z =
0.15) to c. 60% (with z = 0.40) of the species present.
This is often reduced to the rule-of-thumb that a ten-fold
decrease in area leads to a loss of half the species present.
The causes of the species-area relationship appear to be
relatively straightforward, and involve a combination of
sampling effects and environmental heterogeneity. On a
small scale, the increase in number of species with an
increase in area is probably overwhelmingly a result of the
former: that is, put very crudely, a given habitat in which
species are randomly distributed will become increasingly
more completely sampled as the area sampled increases. At
larger scales, environmental heterogeneity will be more
important: that is, as the area sampled increases, so
different habitats with different species in them will be
included in the sample.
Diversity at different scales
Thus the overall diversity of any given area will be a
reflection both of the range of habitats it includes and the
diversity of the component habitats. The greater the
differences between the various component habitats in terms
of species composition, then the greater the overall diversity
will be. The differences between habitats are referred to as
beta (G) diversity, while the diversity within a site or habitat
is alpha (a) diversity. Thus an area with a wide range of
dissimilar habitats will have a high B-diversity, even if each
of its constituent habitats may have low a-diversity.
Differences in site diversity over large areas, such as
continents, are sometimes referred to as gamma (y)
diversity.
An area with relatively low species diversity may therefore
still make an important contribution to the overall diversity
of the larger region it is found in if it contains a significant
number of species which do not occur elsewhere
(endemics). Oceanic islands (see Chapter 14) and
continental montane regions are examples of geographical
entities which typically have comparatively low species
diversity but high rates of endemism.
43
Species Diversity: An Introduction
Assessing the relative importance of areas with high species
diversity and low rates of endemism compared to areas with
lower rates of diversity and high endemism remains an
intractable problem. Attempts have been made to
circumvent this by using somewhat different approaches,
such as Critical Faunal Analyses, but these also generally
do not generate unequivocal results (Chapter 15).
THE GLOBAL DISTRIBUTION OF SPECIES
RICHNESS
Analysis of worldwide trends in biological diversity almost
always treats this in terms of species richness, as this is the
only indicator of diversity for which anything approaching
adequate data is available on a global scale. Biological
diversity is not evenly distributed around the globe.
Latitudinal gradients
The single most obvious pattern in the global distribution of
species is that overall species richness increases with
decreasing latitude. At its crudest this means that there are
far more species per unit area and in total in the tropics
than there are in temperate regions and far more species in
temperate regions than there are in polar regions.
Not only does this apply as an overall general rule, it also
holds within the great majority of higher taxa (at order level
or higher), and within most equivalent habitats, although the
most obvious and frequently cited are forests and shallow-
water marine benthic communities, with, respectively,
tropical moist forests and coral reefs being renowned for
their remarkably high levels of species diversity.
This overall pattern masks a large number of minor trends
where species richness in particular taxonomic groups or in
particular habitats may show no significant latitudinal
variation, or may actually decrease with decreasing latitude;
nevertheless it remains a phenomenon of overwhelming
biogeographical importance.
As well as latitude, changes in diversity can also be
correlated with a many other variables, some of which are
discussed briefly below. For some of these it is not easy to
establish a significant relationship because there are often
confounding variables, and because there are too few
comparable datasets.
The maps in Fig. 5.4 demonstrate broad gradients in
species richness in frogs (left) and trees (right) in the
Americas (data extracted from Duellman, 1988 and Gentry,
1988). For these groups in this part of the world, climatic
factors appear to play a large part in determining such
gradients.
Elevational gradients
In terrestrial ecosystems, diversity generally decreases with
increasing altitude. This phenomenon is most apparent at
extremes of altitude, with highest regions at all latitudes
having very low species diversity. There are fewer
examples showing gradients of species richness with
altitude, although amongst vertebrates this has been
demonstrated for bird species in New Guinea (Kikkawa and
1. Biological Diversity
Figure 5.4 Gradients in species richness: frogs and trees in the Americas
a Sw CE
DA CELTS
repoeibaisiepimsalai
4+ |
Se Bo er ee ee
sejoeds jo seqUINN,
Trees
ay ae
Frogs
a
(data from Dueliman, 1988 and Gentry, 1988)
sejoeds jo JequiNN
Williams, 1971 cited in Brown) and.on the Amazonian
slope of the Andes in Peru (Terborgh, 1977). Gentry (1988)
demonstrates it for woody plants in tropical forests,
although notes that the data for upland sites are very
incomplete. Suggestions have been made that, in tropical
forests at least, diversity may be higher at mid-altitudes
than in lower areas. However, there appear to be no
substantiating data for this ‘mid-altitude bulge’ as a general
phenomenon, although it has been noted in particular cases
such as a desert mountain in Arizona where diversity at
lower and higher altitudes is believed limited by aridity and
low temperate respectively (Brown, 1988).
The decrease in straightforward species numbers with
increasing altitude may in part be a reflection of species-
area relationships, as available area generally decreases with
increasing altitude, and number of species is closely related
to area. Measurement of species numbers in standard-sized
plots, such as those of Gentry (1988) take account of this,
demonstrating that the relationship between altitude and
species diversity is real, although not necessarily
discounting the role that decreased available area may play
in causing this phenomenon. It should also be noted that 8
diversity will often be higher in areas of varied topography
because of increased environmental heterogeneity.
Precipitation gradients
Precipitation is generally believed to be an important factor
governing terrestrial diversity. However, the relationship
between precipitation and diversity is not straightforward,
and it seems that seasonality in precipitation may be as
important as absolute amount. As with altitude, the
relationship between precipitation and diversity is most
apparent at one extreme, as highly arid environments are
well-known to be much less diverse than less arid, or more
mesic, environments at similar altitudes and latitudes. There
are, however, apparently few quantifiable data to
demonstrate this. Gentry (1988) in his study of forest
diversity, demonstrated a strong correlation between plant
species richness and absolute annual precipitation.
However, he notes that this correlation may not apply at all
in the Palaeotropics, and that there were strong indications
that the length and severity of the dry season were more
important than absolute annual rainfall. In the Neotropics,
there is a strong relationship between annual rainfall and
strength of the dry season, which is much less marked in
the Palaeotropics. The importance of seasonality was borne
out by a preliminary study of a Brazilian site with a
relatively low, evenly-distributed annual rainfall, which
showed a much higher species diversity than would be
expected from total rainfall measures alone. Moreover,
there appears to be a marked tailing-off of increasing
diversity with increasing rainfall at high rainfalls, with little
or no increase in diversity once rainfall exceeded 4,000-
4,500mm per year.
However, it should be noted that the limits on diversity may
in fact represent a limitation of sampling technique: in the
two most diverse sites sampled (in areas of year-round
rainfall of 3,000-4,000mm p.a.), diversity was so high in
the plots sampled (in one site 300 species => 10cm diameter
out of 606 individual plants in one plot), that it seems likely
that only by increasing the size of the survey plots would
45
Species Diversity: An Introduction
any further trends be discerned (Gentry, 1988).
Nutrient levels
Although there are few studies of global trends in diversity
and soil nutrients, the relationship between plant community
richness and tropical soil nutrient levels has been the subject
of considerable interest. The data that are available indicate
that the relationship may not be straightforward. Studies in
Southeast Asia indicate that diversity may be highest at
intermediary levels of nutrition, with a decrease at higher
levels, while in the Neotropics diversity generally seems to
increase with increasing nutrient levels, being most strongly
correlated with Potassium (K) levels. This overall trend is
apparently also shown by a variety of other organisms,
including bats, birds and butterflies. In general, however,
diversity in tropical forest ecosystems seems much less
strongly dependent on nutrient levels than other factors,
notably latitude, altitude and precipitation (Gentry, 1988).
The relationship between nutrient levels and diversity in
other ecosystems is also complex: declines in diversity with
increasing nutrient levels of temperate freshwater habitats
(eutrophication) and grasslands are well-documented, but it
is difficult to draw general conclusions from these (Brown,
1988).
Salinity gradients
In aquatic ecosystems, salinity appears to act as a strong
‘normalising’ factor on diversity. Thus, in coastal areas,
diversity almost invariably declines when salinity deviates
from ‘normal’ sea water (i.e. 35 ppt), while in freshwaters
diversity decreases when salinity increases above c. 2 ppt;
this results in a bimodal distribution of diversity with
increasing salinity (Brown, 1988).
Islands
The study of diversity on islands, both real and theoretical,
has been an important factor in much of biogeography and
conservation biology. In particular the equilibrium theories
of island biogeography elaborated by MacArthur and
Wilson (1967) have had an important influence on both
disciplines. More recently discussion in this, as in many
other areas of ecology, has tended to move away from
assumptions of equilibrium to more realistic, but far more
complex, non-equilibrium theories.
SPECIES AND ENERGY
The relationship between diversity and productivity has
been the subject of long-standing debate in ecology. Recent
studies have indicated that available energy is strongly
correlated with species diversity on a large-scale, at least in
terrestrial ecosystems. A study of North American tree
species (Currie and Paquin, 1987) demonstrated that
realised annual evapotranspiration, a measure of available
energy, statistically explained 76% of the variation in
species richness across the continent. Such recent studies
have shown that diversity gradients in tree species are more
closely related to indices of climatic productivity than to
other geographical parameters, including latitude (Adams,
1989). These results could be used to predict accurately tree
1. Biological Diversity
species richness patterns in Great Britain and Ireland.
Preliminary analysis of the diversity of terrestrial
vertebrates in North America apparently yielded very
similar results.
EXPLANATIONS AND HYPOTHESES
The explanation of geographic and temporal variation in
species diversity is one of the central problems of biology.
It has also proved one of the most intractable. The problem
has generated an enormous amount of literature in which
many different hypotheses have been proposed to attempt to
account for it; these hypotheses often operate at different
levels of explanation and much confusion has arisen as a
result. It is beyond the scope of this report to attempt a
thorough review of the subject, although, ultimately, an
understanding of the importance of biological diversity
should rest on an understanding of how and why it has the
form that it does.
It is self-evident that, ultimately, all non-random patterns in
species diversity must depend on past or present variations
in the physical environment. How such variations result in
the patterns observed is often far from clear. It is evident,
however, that any complete explanation must involve both
historical events and current ecological processes - the
former implicit in any explanation of the origin of diversity,
the latter in explanations of its maintenance, these being
46
two separate, although intimately linked, problems. The
relative importance of these two factors in determining
present patterns is still a subject of considerable debate.
References
Adams, J.M. 1989. Species diversity and productivity of trees. Plants
today Nov.-Dec. 183-187.
Brown, J.H. 1988. Species diversity. In: Myers, A.A. and Gillet, P.S.
(Eds), Analytical Biogeography. Chapman and Hall, London.
Currie, D.J. and Paquin, V. 1987. Large-scale biogeographical
patterns of species richness of trees. Nature 329:326-327.
Duellman, W.E. 1988. Patterns of species diversity in anuran
amphibians in the American tropcis. Annals of the Missouri
Botanical Garden 75:70-104.
Gentry, A.H. 1988. Changes in plant community diversity and floristic
composition of environmental and geographical gradients. Annals
of the Missouri Botanical Garden 75:1-34.
Kikkawa, J. and Williams, E.E. 1971. Altitudinal distribution of land
birds in New Guinea. Search 2:64-69.
MacArthur, R.H. and Wilson, E.O. 1967. The Theory of Island
Biogeography. Princeton University Press, Princeton.
Signor, P.W. 1990. The geological history of diversity. Annual Review
of Ecology and Systematics 21:509-539.
Terborgh, J. 1977. Bird species diversity on an Andean elevational
gradient. Ecology 58:1007-1019.
Vidal, G. and Knoll, A.H. 1982. Radiations and extinctions of
plankton in the late Proterozoic and early Cambrian. Nature
297:57-60.
Chapter contributed by Martin Jenkins.
6. MICROORGANISMS
TAXONOMIC SCOPE
This section provides an overview of the phylogenetically
extremely diverse groups collectively regarded as
‘microorganisms’. This term is misleading as by no means
all are microscopic. The definition accepted here is:
organisms which either belong to phyla many members of
which cannot be seen by the unaided eye, or where
microscopic examination, and in many cases growth in pure
culture, is essential for identification (Hawksworth, 1992).
Some of the themes touched on here with specific reference
to microorganisms are developed from a_ broader
perspective elsewhere in the report (Chapter 4). The
glossary should be consulted for definitions of certain
terms.
The classification of the various microorganism groups at
the rank of kingdom, and both below and above that level,
is currently in a state of flux. For the purposes of this
contribution, the terms algae, bacteria, fungi, protozoa, and
viruses are treated in their traditional non-phylogenetic
sense, with some minor modifications (Table 6.1).
However, as the macroalgae (charophytes and seaweeds)
and the lichen-forming fungi (lichens) are discussed
elsewhere in this publication (Chapter 7), these
non-taxonomic groupings are given only brief mention here.
ASSESSMENT OF DIVERSITY
The diversity of microorganisms in terms of the numbers of
species currently known, and those estimated to occur in the
world, was considered by leading specialists in the various
groups at an IUBS/IUMS workshop in 1991 (see below;
Hawksworth and Colwell, 1992 and in prep.). While the
total number of known species is reliably estimated at
159,000 (Table 6.1), considerable difficulty arises in the
estimation of those which remain undescribed.
Nevertheless, the conclusion that less than 5%, and
probably less than 3%, of the world’s microorganisms have
been described is not expected to be unduly pessimistic.
Algae
While the number of recognised algal species can be
asserted with some confidence, the estimated world figure
of 350,000 now proposed has large error margins - indeed
it has been hinted that the chromophyte algae alone might
eventually prove to comprise either 100,000 or up to 10
million species, the diatoms being the most speciose
(Andersen, in press). The terrestrial algal species,
especially those on bark and rocks, and minute ocean
species have received particularly scant attention. Further,
the marine picoplankton, which can make up to 25% of the
phytoplankton biomass in polar waters, were first
recognised only in 1980.
Bacteria
The number of bacterial species accepted in the Approved
List of Bacterial Names was 3,058 in July 1991 (Triiper,
1992); the figure of 4,000 in Table 6.1 has been increased
to allow for cyanobacteria. Perceptions of the true number
of bacteria in the world have changed dramatically during
the last 5-10 years. It has become increasingly evident,
47
Microorganisms
primarily as a result of the application of molecular
techniques (Liesack and Stackebrandt, 1992), that there are
enormous numbers of as yet uncultured bacteria to be found
in soils, deep sea sediments, as mutualists in protozoans and
other organisms and, most importantly, in the digestive
tracts and pockets of a wide variety of animals - including
most insects (Triiper, in press). It has been suggested that
one genus of wall-less bacteria inhabiting insect guts, the
mollicute Spiroplasma, may prove to be the largest genus
on Earth with well over one million species (Whitcomb and
Hackett, 1989).
Fungi
The number of fungi estimated to occur in the world has
recently been conservatively estimated at 1.5 million species
(Hawksworth, 1991a). This figure contrasts markedly with
the 70,000 now described - that figure has been increased
from the 69,000 cited by Hawksworth (op. cit.) to allow for
fungi newly published since 1990. The 1.5 million figure is
conservative as in the calculations leading to it: (1) a
modest world estimate of vascular plants was employed, (2)
no special allowance was made for fungi to be expected on
the large numbers of insects now postulated, (3) the UK
vascular plant:fungus ratio of 1:6 used must be an
underestimate as additional fungi continue to be found in
that country, and (4) no provision was made for any
proportionately increased numbers in the tropics or polar
regions. Whether an upward revision of the 1.5 million
figure is defensible must await in-depth studies of particular
tropical sites. See Chapter 4 for an alternative view.
Protozoa
Corliss (1991) estimated the number of known non-fossil
protozoan species at 40,000. No calculated predictions of
the number of world species have been prepared, but many
groups, such as the heterotrophic heterokonts in soil, have
scarcely been investigated. The total world estimated
number of 100,000 used here could prove to be a gross
understatement.
Viruses
No comprehensive catalogue of the world’s known viruses
currently exists, but it is expected that about 5,000 will be
recognised in a compilation being planned by the
International Committee on the Taxonomy of Viruses for
publication in 1993. The estimate of 500,000 species
presented here reflects the substantial numbers of new
viruses to be expected on yet unstudied non-crop plants, and
especially insects. Also scarcely investigated are viruses
only recently recognised as frequent in marine plankton,
plasmids in fungi, and phages on bacteria - not least on
‘unculturable’ bacteria.
SPECIES CONCEPTS IN MICROORGANISMS
Comparisons of species numbers between microorganisms
and macroorganisms, and indeed also between the different
microorganism groups, are complicated by variations in
species concepts. While the idea of the ‘biological species’
is not without appeal to microbiologists, in practice in the
majority of cases it is not readily applicable. This difficulty
arises both because sexual processes are absent or difficult
1. Biological Diversity
Table 6.1 Estimates of the number of described species and possible
undescribed species of microorganisms
GROUP DESCRIBED ESTIMATED PER CENT
SPECIES SPECIES KNOWN
Algae 40,000 350,000 11.0
Bacteria (incl. cyanobacteria and 4,000 3,000,000 0.1
‘unculturables’)
Fungi (incl. yeasts, lichen-forming 70,000 1,500,000 5.0
fungi, slime moulds, and
oomycetes)
Protozoa (proctoctists, excl. algae 40,000 100,000 40.0
and oomycete fungi) :
Viruses (incl. plasmids, phages, 5,000 500,000 1.0
etc.)
TOTAL 159,000 5,450,000 3.0
Source: Based primarily on data in Hawksworth, D.L. and Colwell, R.R., (Eds) (in prep.). Biodiversity amongst microorganisms and its
significance. Biodiversity and Conservation 1.
to detect in many microorganism groups, and further when
they are known to occur it is often impractical to determine
breeding groups. In practice microbiologists tend to be
pragmatic, recognising as ‘species’ specimens or strains
with a high degree of morphological, biochemical, or
molecular similarity and which produce replicating lineages.
The scale of characters used is inversely proportionate to
the size and number of morphological characters. In the
bacteria and yeasts suites of assimilation, substrate
utilisation, and cultural attributes are extensively used,
while these feature to a much lesser extent in algae,
filamentous fungi, and protozoa.
Stress is invariably placed on the recognition of marked
discontinuities in several characters, but emphasising those
features which are relevant in human terms - for example
the ability to cause diseases in particular animals or plants,
to form toxins, to conduct economically important
fermentations, or to produce desired chemical products.
Clones, the progeny derived from a single cell and which
do not exhibit any genomic variation or recombination, are
to be found in all microorganism groups. However, these
are not always easy to recognise, and the practice has been
to accept as species clones fulfilling the requirements of
distinctness normally associated with that rank. Clones are
frequently opportunistic organisms well-adapted to
particular ecological niches; in the case of the conidial
fungi, clones are derived from a part of the life-cycle of
sexually reproducing species in which the sexual stage has
sometimes been entirely lost.
Particular aspects of the use of species concepts in the
different microorganism groups are considered further
below:
Algae
The biological species concept is theoretically usable in
those algae which are entirely sexual or have such stages in
their life cycles, but experimental verification is rarely
practical as many species cannot be readily grown in pure
48
culture. Its application in practice has thus been extremely
limited. In large groups such as the diatoms and coccoliths,
while sexual stages are known or expected to occur, in
reality morphological species concepts have to be used,
increasingly employing characters only visible by the
Scanning Electron Microscope (SEM). In eight algal
classes, however, sexual reproduction is entirely unknown.
Chemical characteristics are extensively used as aids to
species differentiation in certain groups (Kessler, 1985).
Mating complex studies and molecular approaches are also
increasingly being used. The latter approaches are
illustrating that considerably diverse taxa are sometimes
grouped in the same genus or species, although the
converse situation is also known. An overview of species
concepts in algae is provided in Andersen (1992).
Bacteria
As sexual differentiation does not occur in bacteria and
most reproduction is asexual, and further, as recombination
between different strains is difficult to detect, species
concepts in bacteria have largely been based on overall
similarities. Since the early 1960s, numerical taxonomic
studies utilising 50 to several hundred biochemical and
cultural tests have played a major role in defining bacterial
species. Similarity coefficients are computed, and phenetic
groups formed at about the 80-85% similarity level are
generally taken as equivalent to species (Austin and Priest,
1986; Sneath, 1989). The advent of molecular techniques
has enabled species concepts derived from phenetic methods
to be reassessed. DNA homologies of 20-50% are found
between species in the same genus, and 60-70% between
subspecies within the same species (Johnson, 1989). The
International Commission on Systematic Bacteriology
recommends that a minimal DNA homology of 70% be
required for species-level treatment (Wayne, 1987).
Fungi
A consequence of the wealth of morphological characters in
fungi is that species continue to be mostly distinguished by
marked discontinuities between those features. The
assimilative and predominant phase is haploid, and most
fungi are either sexual or derived from ancestors that were
so. Despite the considerable literature on speciation in fungi
(Burnett, 1983), the delimitation of populations from a
biological standpoint remains in its infancy. As particular
examples are studied in depth, it is becoming increasingly
apparent that several discrete reproductively isolated groups
are not uncommonly present within single morphospecies
(Brasier, 1986). Mycologists have been reluctant to
recognise such groups at the rank of species, but this can be
expected to change where particular groups also have other
important features such as pathogenicity to different crops.
While a wide range of biochemical and molecular
techniques are currently being employed in the fungi
(Hawksworth and Bridge, 1988), the application of many of
these is limited to the 20% of the known species which can
be grown in pure culture. Where DNA homology studies
have been conducted, notably in yeasts and certain
economically significant genera such as Aspergillus, the
differences between morphological species tend to be in the
20-50% range (Kurtzman, 1985), as they are in bacteria.
Protozoa
In contradistinction to the fungi, many protozoan species
are diploid. In numerous groups information on life-cycles
and sexuality are still lacking, rendering it difficult to apply
a biological species concept. Clonal protozoans are,
however, often described as species, while in contrast, as
in the case of fungi, morphologically defined species may
be found on more critical analysis to consist of a number of
discrete gene pools.
Viruses
While some biologists are reluctant to recognise viruses as
‘living’, that they are functional biological entities is
inescapable. They possess genomes, replicate, evolve,
occupy specific ecological niches, and exhibit intrinsic
variability. Ultrastructure, serological tests, physical and
chemical structure and features, and the ability to infect
particular hosts are used in species separation. The species
concept in virology has been analysed by Regenmortel
(1990). He took a pragmatic stance and defined a virus
species as a polythetic class of viruses constituting a
replicating lineage and occupying a particular ecological
niche. This definition has the attraction of being applicable
both to groups which are able to undergo recombination and
those which are clonal.
EXTENT OF GENETIC DIVERSITY
The extent of genetic diversity exhibited by microorganism
groups is vast in comparison to that of macroorganisms.
This conclusion was to be expected bearing in mind that the
earliest bacteria probably arose around 3.5 billion years ago
on an Earth formed only one billion years earlier, whereas
the first land plants, for example, did not emerge until
about 0.4 billion years ago; i.e. microorganisms have had
nine times as long to diverge as land plants.
This diversity is illustrated to some extent in terms of the
numbers of phyla recognised, but most forcefully at the
molecular level. Of the 95 phyla accepted by Margulis and
Schwartz (1988), 52 belong to the microorganisms as
defined here (/ess the virus groups not considered by those
authors). More significantly, the study of 16S-like rRNAs
49
Microorganisms
in prokaryotes led to the suggestion that they should be split
into two separate groups, Archaebacteria and Eubacteria,
and that these were roughly equivalent to the Eukaryotes.
Recognising that most biologists would be unwilling to
accept plants and animals as belonging to the same
kingdom, the higher rank of "domain" has been applied to
these three groups, i.e. the domains Archaea, Bacteria, and
Eucarya (Woese ef al., 1990). Studies with the gut
protozoan Giardia lamblia, however, have further
demonstrated that at least some eukaryotic microorganisms
are much more remote from each other than had hitherto
been assumed; for example, on the basis of 16S-like
tRNAs, the crustacean Artemia salina and Homo sapiens
are ten times closer to each other than either are to Giardia
(Sogin, 1991).
The extent of genetic diversity now demonstrated between
the higher ranks of microorganisms is reflected also at the
species level. Both the genetic diversity within single
microbial species, and that between several species referred
to the same genus, can also be vast in comparison with
macroorganism groups. This is especially true at the DNA
homology level where 20-50% similarities are regularly
encountered between species (see above), whereas primate
‘species’ may still be regarded as distinct although sharing
90+ % DNA homology.
One consequence of the considerable genetic diversity
within microbial species is that in certain microorganism
groups infraspecific categories are utilised to an extent not
otherwise seen outside the higher vertebrates. These include
subspecies, pathovars, ‘special forms’, and serotypes. In
addition, complex race notations have been developed
within particular species of major medical or plant
pathogenic importance. This tradition has developed as a
pragmatic response to the need to label populations to a
finer degree because of the different effects they have on
humans or their crops.
From this discussion it will be apparent that if identical
DNA homology criteria were used for species separations
in both macro- and microorganisms, the numbers of known
and estimated species in Table 6.1 would have to be inflated
by not less than an order of magnitude.
REGIONS AND HABITATS OF MAXIMUM
DIVERSITY
The variety of ecological niches exploited by the major
groups of macro- and microorganisms is directly related to
their geological age; ecology recapitulates phylogeny (Price,
1988). The greatest niche breadth is consequently seen in
the bacteria, and then, in declining sequence, in the algae
and protozoa, fungi, animals, and plants.
While there is every reason to suppose that regions and
habitats with a maximum diversity of macroorganisms will
also be particularly rich in microorganisms - a consequence
of the larger numbers of host-specific parasites, mutualists,
and saprobes to be expected - there are additional habitats
of no importance for macroorganisms which are important
for the conservation of microorganism diversity.
Amongst the bacteria are species able to grow in extreme
1. Biological Diversity
saline substrata or at high sugar (low water activity)
concentrations, ones which thrive at high concentrations of
heavy metals, sulphur, or other generally toxic compounds,
major groups restricted to anaerobic situations, and ones
able to tolerate or even thrive at extremely high (e.g.
Thermotoga lives at 90°C) or low temperatures (e.g. at or
below freezing point in the Antarctic).
Triiper (in press) identified the following environments as
ones dominated by microorganisms or ones which are
strongly influenced or stabilised by them:
© hypersaline neutral and alkaline lakes (salt lakes and
soda lakes), e.g. East African rift valley lakes, the Dead
Sea
e hot springs (hydrotherms, fumaroles, solfatoras) which
have not been disturbed
e natural leaching environments (acid crater lakes, acid
mine waters)
® peat mosses, permafrost tundra, cypress and mangrove
swamps
© stratified (meromictic) lakes
e@ hot deserts (sand and rocks) which have not been
disturbed)
@ bare lichen-encrusted rock areas (with associated bacteria
and fungi), in all climatic regions
© estuaries (salt marshes, mud flats, beaches)
e deep sea environments (hydrothermal vents, hypothermal
zones, Manganese nodule areas).
Extreme environments also continue to be a particularly rich
source of previously unknown microorganisms belonging to
diverse groups. Even though not all the species are known,
it is evident that due accord needs to be given to extreme
environments when drawing up international, national, or
regional plans for the establishment of protected areas.
As a consequence of the antiquity of the groups, there is a
tendency for microorganisms to have much broader
geographic ranges than macroorganisms. Biogeographic
studies, except in the case of lichens and macroalgae, are
rarely undertaken. However, there is no reason to suppose
that while there are a considerable number of almost
cosmopolitan species, many others do not have
geographically restricted ranges. This is certainly true for
the fungi, but current perceptions of distributions on a
global scale are skewed by inadequate sampling.
Mycologists, for example, would take in their stride the
discovery of a species previously known only from Europe
in an undisturbed habitat in Australia, whereas a similar
event would cause amazement among workers in most other
groups.
Conversely, detailed biogeographic analyses from the world
level down to national mapping programmes, clearly
demonstrate that in the fungi numerous species are narrowly
restricted geographically. Studies on the numbers of species
of particular families and genera of fungi in different
geographic regions can potentially lead to the recognition of
centres of maximum diversity, as demonstrated for certain
ascomycete groups by Pirozynski and Weresub (1979). A
shortage of authoritative inventories and surveys currently
50
limits the utilisation of such approaches in site-selection.
ROLE OF MICROORGANISMS IN BIODIVERSITY
MAINTENANCE
Microorganisms have played a major role in the evolution
and diversification of macroorganisms. They contributed
key organelles such as mitochondria and chloroplasts to
eukaryotic cells, and as mutualists are either involved in
nutrient-supply or perform other biochemical processes on
which they depend (Margulis and Fester, 1991). Bacteria,
fungi, and protozoa in the guts of insects and herbivorous
mammals perform crucial roles in their digestive processes,
particularly in the breakdown of celluloses and lignins, and
without which they could not exist (Smith and Douglas,
1987). About 85% of the Earth’s vascular plants form
mycorrhizas with fungi. This life-style is often obligate in
nature, the mycorrhizas being crucial to the absorption of
growth-limiting nutrients (Read, 1991). The very existence
of many macroorganisms is consequently dependent on the
continued availability of the mutualistic microorganisms
they require.
In the marine environment, up to 80% of the biomass and
productivity in open waters is contributed by ultraplanktonic
algae (Andersen, 1992). Further, dinoflagellates form
mutualisms with coelenterate stony corals, and the outer
ridges of major reefs taking the full force of the oceans are
formed by crustose coralline algae cementing detritus
together (Round, 1981). In the absence of these mutualistic
microorganisms, coral reef ecosystems simply could not
exist (Smith and Douglas, 1987). Without the coral
mutualists one of the most biologically diverse habitats on
Earth would never have been formed.
At the ecosystem function level, food networks of all life on
Earth are ultimately dependent on microorganisms. This
holds for terrestrial and marine ecosystems (Andersen,
1992; Grassle et al., 1991; Price, 1988), yet ecologists and
conservationists only exceptionally take it into account.
The greatest biomass in soil, on the basis of current
evidence, is that of the microorganisms, especially the fungi
(Lee, 1991; Lynch and Hobbie, 1988). These play a variety
of roles related to the maintenance of soil structure and
composition both through the biodegradation and
incorporation of dead plant and animal remains, and by
extra-cellular fungal polysaccharides which bind soil
particles together, thus increasing soil aggregation and
stability (Lal, 1991).
Microorganisms also contribute to the maintenance of
ecosystem structure through natural biocontrol. Plant
pathogenic microorganisms can limit plants that would
otherwise expand explosively in the absence of their
co-existing pathogens. Similarly, entomogenous
microorganisms can limit the populations of insects that
would otherwise become major pests (e.g. defoliants) of
trees or other plants. In these two cases, if their targets
have crucial ecological roles, the loss of the containing
microorganism would lead to major changes in the
ecosystem.
ROLE OF MICROORGANISMS IN BIOSPHERE
FUNCTIONS
Bacteria shaped the early atmosphere of Earth, the start of
life coinciding with a fall in carbon dioxide and an increase
in methane at around 3.8 billion years ago. The
photosynthetic cyanobacteriawere subsequently instrumental
in producing oxygen, and microorganisms on land would
have increasingly removed carbon dioxide from the early
atmosphere in rock weathering (Lovelock, 1988). In the
absence of these activities there would have been no
macroorganisms or humans. Microorganisms continue to
play a major role in the maintenance of the biosphere and
global ecology through the various biogeochemical cycles.
They perform unique and indispensable roles in the
circulation of matter in the world (Stolz et al., 1989). The
principal biogeochemical cycles with which they are
involved are:
Carbon
It has been estimated that about 40% of the carbon fixed by
photosynthesis on the Earth is carried out by algae and
cyanobacteria, especially those in oceans and seas. Bacteria
also fix atmospheric carbon dioxide anaerobically and in
methanogenesis. Methanogenic archaean bacteria generate
about 58 % of the Earth’s methane. Conversely, wood-decay
fungi are instrumental in releasing around 85 billion tonnes
of carbon (as carbon dioxide) into the atmosphere each
year. Ruminant gut microbial populations also produce
methane, and other methyl gases are produced by fungi
during wood decay. The tissues of microorganisms further
have roles as carbon sinks, and their removal of carbon
from the atmosphere in rock weathering is an on-going
process.
Nitrogen
The Earth’s nitrogen cycle is dependent on bacteria
(including cyanobacteria) for nitrogen fixation, the oxidation
of ammonia, nitrification, and nitrate reduction. The
magnitude of the amounts involved is staggering: each year
bacteria fix 240 Tg of nitrogen, release 210 Tg of nitrogen
by denitrification, and release 75 Tg of ammonia (Triiper,
1992).
Sulphur
The sulphur cycle on Earth is dependent on sulphur-
reducing bacteria for the reductign of sulphate into
hydrogen sulphide, on purple and green photosynthetic
bacteria for the oxidation of sulphides to sulphur, and
sulphur oxidising bacteria for the conversion of sulphur to
sulphates. Bacteria are also involved in the biogenesis of
dimethylsulphide, a substance of particular relevance as a
greenhouse gas and postulated as performing an
equilibrating function for the planet (Lovelock, 1988).
Minerals
Microorganisms of various types, including algae, bacteria,
fungi, and protozoa, are important in the production of a
wide range of biogenic minerals, notably in the processes
of rock weathering. These include diverse kinds of
carbonates, phosphates, oxalates, sulphates, silicates,
sulphides, and further oxides of iron and manganese
(Krumbein, 1983; Leadbeater and Riding, 1986; Stolz et
al., 1989).
51
Microorganisms
POTENTIAL CONTRIBUTION OF
MICROORGANISMS TO SUSTAINABLE
DEVELOPMENT
Microorganisms have the potential to contribute to
sustainable developmentin multifarious ways (Hawksworth,
19916; Persley, 1990). Production on existing agricultural
land may be increased through:
e the selection and introduction of the most efficacious
nitrogen-fixing Rhizobium strains into legume crops
e@ the enhancement of natural nitrogen fixation by the
application of cyanobacterial inocula, either directly or
through mutualists (e.g. improvement of cyanobacteria
of Azolla for use in rice-fields)
® the use of bacteria and fungi as biocontrol agents for
insect pests, plant pathogens, disease vectors, and
noxious weeds
© the mass production of the most efficacious mycorrhizal
(and in the future almost certainly also beneficial
endophytic) fungi for inoculation into seeds or seedlings
on or prior to planting.
Genes from bacteria and fungi with useful properties, for
example the production of an insecticidal metabolite or
enzyme, can be cloned and inserted into the genome of a
crop plant by an increasing range of methods. Indeed, a
plasmid in the crown gall bacterium Agrobacterium
tumefaciens is well-established as a practical mechanism by
which genes from any source can now be engineered into
over 20 major world crops.
A wide array of pharmaceutical and other industrial
products are already obtained from microorganisms grown
under factory conditions. These include, for example,
organic acids, vitamins, antibiotics, anti-inflammatory
drugs, immunoregulators (e.g. cyclosporin from a saprobic
fungus which is now routinely used in human transplant
surgery), food colourings, fragrances, and food
preservatives. The discovery and studies of the actions of
naturally occurring compounds can also lead to
semi-synthetic drugs of great potential, as in the case of
ivermectin first used against helminths parasitic on livestock
but now also employed in humans against onchocerciasis
(river blindness).
In addition, cellulosic and lignosic wastes from agricultural
and industrial sources can be biodegraded by
microorganisms and converted to animal feedstuffs.
Waste-water treatments using anaerobic bacteria and
filter-feeding ciliates reduce pressure on freshwater
supplies. Microorganisms are crucial to the functioning of
sewage filter-beds. Bacteria can also be employed in the
removal of toxic chemicals, especially heavy metals, from
liquid waste; any valuable metals can be recovered for
reuse. The bioremediation of major oil spills at sea can be
achieved by applying nitrogen fertilizers which encourage
the naturally presenthydrocarbondegrading microorganisms
to proliferate.
Biogas (methane) production from a variety of agricultural
and other wastes for use as fuel is dependent on anaerobic
bacteria. This has the potential to reduce the pressure on
1. Biological Diversity
forests by providing an alternative energy supply.
An expanded range of sources of food for humans can be
derived both from the mass-production of certain algae and
filamentous fungi (e.g. the Fusarium graminearum strain in
‘Quorn’), and through the increased use of waste materials
for the commercial production of a wide range of edible
macrofungi.
The design and development of technologies to increase the
utilisation of microorganisms for human benefit therefore
merit interpretation as activities integral to the formulation
of long-term sustainable development programmes.
THE NEED FOR DIVERSITY AMONGST
MICROORGANISMS
While sufficient diversity of microorganisms to enable the
various functions necessary for ecosystem maintenance and
the operation of biogeochemical cycles is clearly crucial,
the extent to which individual species are important is less
certain. In monitoring microorganisms with reference to the
conservation of biodiversity in macroorganisms, the
maintenance of functional groups rather than individual
species can be presumed to be limiting - except where a
particular microorganism is a keystone species.
There has been considerable debate as to the significance of
functional redundancy in ecosystem function and
maintenance (Solbrig, 1991). The presence of a wide
variety of species able to perform similar roles is
unquestionably beneficial as it provides an ecosystem with
increased resilience to perturbations. For example, in the
case of ectomycorrhizas of temperate and boreal forests, the
ability of a tree to form associations with a variety of fungi
(over 100 in the case of Berula) enables that tree to grow
satisfactorily even if only a few of the candidate
mycorrhizal fungi are present in a particular soil. Further,
if the mycorrhizal species are differentially sensitive to
pollutants, the tree can continue provided at least some of
those fungi can tolerate the ambient pollution levels. In the
event that too many species from a functional group are
eliminated, at some point an ecosystem will start to break
down irretrievably. In.this regard, the implications for trees
of the recently reported widespread and dramatic losses of
ectomycorrhizal fungi in Europe are of particular concern
GJaenike, 1991).
Single microorganisms can also function as keystone species
crucial to the maintenance of particular ecosystems. This
applies to marine environments such as coral reefs, kelp
forests formed by Macrocystis in temperate waters, and
lichen-dominated deserts, heaths and rocks. Microorganisms
are most important as keystone organisms when they
function as mutualistic symbionts in organisms that
dominate an ecosystem, and in low productivity/high
diversity systems (Solbrig, 1991). Examples include
dinoflagellates in corals, endomycorrhizal fungi in tropical
forest trees, and nitrogen-fixing bacteria in tree roots.
Individual microorganisms which are major parasites can
also function as keystone species through natural biocontrol
processes. For instance, trypanosomes in East Africa keep
cattle out of wide areas and so may limit soil degradation.
52
The present state of ignorance of the biology, ecology, and
biochemical activities of so many microorganisms is
comparable to that of their role in food-webs (cf. above). It
is consequently often difficult or impossible to assert
whether a particular microorganism is functionally
redundant or a keystone species. Thus, while the presence
of a variety of lignosic wood decay fungi might at first be
assumed to be a case of functional redundancy, in practice
the species of wood attacked can be restricted, and in most
instances the specific enzymes being formed are unknown.
Several species of fungi with different but complementary
properties may need to work simultaneously or
successionally in the decay of a single log. Furthermore,
one or more of the decay fungi in that log might be a
source of digestive enzymes for an insect of ecological
importance in that ecosystem (Martin, 1987).
EX SITU CONSERVATION OF MICROORGANISMS
A wide range of techniques is available for the preservation
of microorganism strains, freeze-drying (lyophilisation) and
storage in liquid nitrogen (cryopreservation) being the most
efficacious for long-term storage. Although not all
microorganisms can yet be preserved by such methods, the
development of programmable coolers and cryomicroscopy
is enabling protocols to be devised for the successful
cryopreservation of organisms previously considered
recalcitrant. Even where species cannot be grown in pure
culture, host tissue including them (e.g. plant leaves
infected with rust fungi) or samples of the substrate itself
(e.g. soil) can be conserved by cryopreservation. A survey
of the existing technology is provided by Kirsop and Doyle
(1991), and the World Federation for Culture Collections
(1990) has issued guidelines for the establishment and
operation of such collections.
Further information on ex situ culture collection is provided
in Part 3.
THE TAXONOMIC CHALLENGE
Studies on the biodiversity and roles of almost all
microorganism groups are frustrated by an inadequate
taxonomic base. Not only are there vast numbers of species
yet to be described, there are few modern monographs,
keys, and other readily available aids, and
disproportionately few taxonomists so that assistance with
identifications is difficult to obtain. This issue requires
priority attention at national, regional, and international
levels. It is clearly unrealistic for most countries even to
contemplate the provision of comprehensive microorganism
identification services. However, attention could be focused
on strengthening existing centres of expertise, developing
north-south and south-south linkages, establishing networks
of centres and specialists, and endeavouring to ensure that
research agendas are complementary and collaborative.
Action to improve the knowledge base
An IUBS/SCOPE workshop on Ecosystem Function of
Biological Diversity held in Washington DC in June 1989
recognised that the issue of microbial diversity and its
function had been neglected and was in urgent need of
attention; the workshop recommended that [UBS and IUMS
(International Union of Microbiological Societies) establish
a cooperative programme to address this problem (Di Castri
and Younés, 1990).
An IUBS/IUMS workshop on Biodiversity amongst
Microorganisms and its Relevance was therefore convened
in Amsterdam 7-8 September 1991. Representatives of
relevant international scientific organisations concerned with
different groups of microorganisms, together with other
specialists, presented overviews of the current knowledge
base (Hawksworth and Colwell, in prep.). A 14-point action
statement, MICROBIAL DIVERSITY 21, was drawn up
detailing the remedial work necessary to raise to an
appropriate level our knowledge of the biodiversity of
microorganisms and its relevance. The various action points
are currently being developed and costed, but it must be
recognised that substantial international resources will be
required to implement this programmeat the level necessary
for it to realise its objectives. This Chapter draws heavily
on the presentations and discussions which took place
during the TUBS/IUMS workshop and the proceedings
(Hawksworth and Colwell, in prep.) should be consulted for
further information on many of the topics discussed here.
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Brasier, C.M. 1986. The dynamics of fungal speciation. In: Rayner,
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Corliss, J.O. 1991. Introduction to the protozoa. In: Harrison, F.W.
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Grassle, J.F., Lasserre, P., McIntyre, A.D. and Ray, G.C. 1991.
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Hawksworth, D.L. (in press). Biodiversity in microorganisms and its
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Hawksworth, D.L. and Bridge, P.D. 1988. Recent and future
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Kirsop, B.E. and Doyle, A. (Eds) 1991. Maintenance of
Microorganisms, 2nd edn. Academic Press, London. 308pp.
Krumbein, W.E. (Ed.) 1983. Microbial Geochemistry. Blackwell
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Kurtzman, C.P. 1985. Molecular taxonomy of fungi. In: Bennett, J.W.
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Global Ecology. Academic Press, San Diego. Pp.31-49.
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in
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54
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This section was prepared by Professor D.L. Hawksworth,
International Mycological Institute, UK.
7. LOWER PLANT DIVERSITY
The term ‘lower plants’ is a convenient but imprecise label
for a disparate group of plants and plant-like organisms
which are defined primarily by their lack of vascular tissue
(the transport system for water and nutrients within higher
plants). Under this heading we here discuss bryophytes,
lichens and larger algae. Many authorities would only
include the first of these among the ‘true’ plants (defined as
those developing from an embryo; see Chapter 8). The
lichens are composite organisms, not true plants, discussed
here for convenience.
BRYOPHYTES
The bryophytes comprise some 14,000 species, consisting
of 8,000 mosses and 6,000 liverworts. This is a very
diverse group of plants containing several classes that are
only distantly related. These classes, and their main
subdivisions (orders) vary in their evolutionary history and
geographical points of origin, and hence vary also in their
current regions of maximum abundance and diversity. On
a global scale, therefore, a more accurate assessment of
areas of biodiversity should rely more on numbers of taxa
within major taxonomic divisions of the bryophytes than on
the oversimplified picture derived from crude summations
of the whole group. Nevertheless, as with other plants,
certain areas of the world are recognised as being
particularly rich in bryophyte species, usually (but by no
means always) the same areas where mosses and/or
liverworts form more than 50% of the active biomass. In
general terms, although bryophytes occur almost throughout
the world, the majority of taxa are distributed in areas of
high oceanicity, i.e. with cool or temperate, consistently
moist climates. Their maximum diversity is to be found in
regions where such conditions have persisted over
geological time, and where tectonic factors have brought
about an amalgamation of several regional floras. The
tregions of high species richness are noted in Table 7.1.
In contrast to many groups of mosses, liverworts generally
(with the exception of the highly adapted Marchantiales)
Table 7.1 Regions of high bryophyte
diversity
REGION SPECIES
(approximate)
Indo-Australian archipelago 3,000
(esp. New Guinea, Sulawesi and Borneo)
South America 3,000
(temperate, montane)
S Australasia 2,400
(esp. Tasmania and New Zealand)
N America 2,000
(Pacific, subarctic)
NE Asia 2,000
(Pacific, subarctic)
Himalayas 2,000
E. Africa 2,000
(and adjacent islands)
Europe 1,800
(Atlantic areas, incl. British Isles)
British Isles 1,000
55
Lower Plant Diversity
show little adaptation to desiccation, either physiologically
or by reduction from perennial to annual growth cycles. In
general, therefore, liverworts reach their maximum
diversity and only achieve dominance in highly oceanic
regions. There are fewer recognised genera than in the
mosses but this is offset to a degree by the much larger
numbers of species in some of them (e.g. Frullania, with
up to 400 species; Plagiochila, with about 500). For
convenience, the Hornworts (Anthocerotae) are included
here with the liverworts.
Both mosses and liverworts (and hornworts) consist of
major divisions into orders and families that may have
widely different habitat preferences. There are too many
such divisions to detail here, but the more significant
groups (orders and families) are listed in Tables 7.2 and 7.3
to provide a reasonably representative picture.
LICHENS
Lichens are composite organisms consisting of a usually
dominant fungal partner in symbiosis with one or more
photosynthetic partners, the resulting composite, organised
structures behaving as independent entities. The fungal
partner (mycobiont) is, in most cases, an ascomycete, rarely
a basidiomycete, while the autotrophic partner (photobiont)
may be a green alga or a cyanobacterium. Lichen
photobionts come from a small number of genera most of
which occur widely in nature while lichen mycobionts are
exclusively lichen-forming and are taxonomically diverse,
many coming from orders that also have non lichen forming
taxa. The lichen symbiosis is one of the most successful
known in nature. Of the 46 orders in the Ascomycotina
some 16 have lichenised taxa to a greater or lesser degree,
and out of some 238 families, 81 consist entirely of lichens
or at least have some lichenised taxa. Lichenisation is a
polyphyletic process that has occurred at many different
times.
Currently, the consensus of known lichenised taxa world
wide varies from 13,500 to 17,000. On the basis of recent
monographic revision of a number of widespread lichen
genera, and the collection of lichens from areas of the
world previously unknown or little known lichenologically,
it is safe to assume that a realistic world total for lichens
will be closer to 17,000 and possibly even to 20,000. It
seems probable that at present we know 50-70% of the
world’s lichens, though future discoveries of short-lived,
fast-growing lichens on leaves and on bryophytes, and of
Southern Hemisphere microlichens, could substantially alter
this estimate (Galloway, 1992).
Although for higher plants the tropics are regarded as major
sites of biodiversity, much less is known about tropical
lichens whose biodiversity tends to be richest in canopy
vegetation, which is still very poorly sampled in many
tropical areas. Temperate areas of the world, on the other
hand, with their wide variations of habitat, geology and
climate are known to be major sites of lichen diversity. Of
great importance are the temperate rainforests of the
Southern Hemisphere, especially South America, New
Zealand, Tasmania, south eastern Australia and the
highlands of the tropical Pacific islands.
1. Biological Diversity
Table 7.2 Selected orders and families of mosses
DISTRIBUTION AND ECOLOGY
Cosmopolitan. About 80 species. Maximum diversity in cool oceanic regions of N
Hemisphere: c. 40 spp in W Europe, similar in N America; 13 in SE Asia and Pacific;
15 in tropical S America; 13 in E. Africa; < 6 in Australasia? Terrestrial, mainly
Old and diverse group with regional endemism and widely differing areas of
Cosmopolitan; tropical montane and high latitude; greatest diversity probably in W
Europe, N America and NE Asia. Mainly calcifuge.
Cosmopolitan; most diverse in, Mediterranean or continental climates, extending to
semi-deserts; principally in temperate to subarctic N Hemisphere but strongly
represented in Australasia and Africa. Xerophytic on soil and rocks, rarely epiphytic,
Lowland tropics and subtropics. Greatest diversity in SE Asia and W Pacific; absent
Mostly subarctic and alpine. Highest diversity in W Europe and N America.
Cosmopolitan; most numerous in cool temperate to polar regions; ecologically
Ecologically very important in the epiphytic biome of the montane tropics. Major
diversity in W Pacific (esp. New Guinea) with other areas in Australasia, S America,
Endemic to W Pacific (esp. New Guinea). Epiphytes.
Endemic to W Pacific and Australasia. Epiphytes and lignicoles.
Mainly cool temperate to arctic with high diversity in NW Europe, N America and NE
Tropical montane rainforests, especially abundant in SE Asia and W Pacific where
Tropical montane rainforests, especially SE Asia and W Pacific. Pendulous epiphytes.
Greatest diversity in the humid tropics, especially S America and SE Asia with
Temperate to tropical montane. Maximum diversity in SE Asia/W Pacific (esp.
Indonesia, Papua New Guinea) and tropical America. Mainly acidophilous, lignicolous
Temperate to arctic. Greatest diversity in N America, NW Europe and NE Asia.
FAMILIES GENERA
Sphagnales
1 1
calcifuge.
Polytrichales
2 21
diversity. About 200 species.
Bryales
85 families incl: Most mosses; about 7,000 species.
Dicranaceae 45
Pottiaceae >70
many annuals.
Calymperaceae 12
from cool temperate regions.
Grimmiaceae 12
Bryaceae 20
important in polar deserts.
Orthotrichaceae 21
E Africa. Mainly photophilic epiphytes.
Spiridentaceae 2
Hypnodendraceae 2
Amblystegiaceae 21
Asia. Hygrophilous and subaquatic.
Pterobryaceae 30
many genera endemic. Frondose epiphytes.
Meteoriaceae 19
Hookeriaceae 27
significant endemism in both. Hygrophilous.
Sematophyllaceae 49
and epiphytic.
Brachytheciaceae c 30
Mainly ground-dwelling.
Hypnaceae 40
Cosmopolitan but with strong regional speciation in all of the areas mentioned
above. ’
Other regions of important local, lichen biodiversity are the
unique coastal fog lichen communities (nebeloasen) found
in northern Chile, Peru, Baja California and Namibia,
where members of the family Roccellaceae are particularly
well-developed.
Islands also often show high lichen biodiversity in
comparison with large continental areas, not only islands
surrounded by water, but biogeographical islands (i.e. areas
of habitat or climate diversity such as rock outcrops,
mountains or ranges in an otherwise uniform forest or
56
grassland landscape). Lichens are particularly successful
pioneer colonisers, and so are important components of
vegetation in many harsh environments of the world, such
as alpine and polar regions, in hot and cold deserts, and in
often toxic, mineralised environments.
Comparative figures of lichen diversity for a number of
areas are presented in Table 7.4. The information is derived
from published accounts of varying age and reliability, most
of which are recorded in the bibliography of Hawksworth
and Ahti (1990), and from unpublished data.
Lower Plant Diversity
Table 7.3 Selected orders and families of liverworts (including hornworts)
DISTRIBUTION AND ECOLOGY
Living fossils with disjunct distributions, all in moist temperate habitats.
An ancient group best represented in the W Pacific.
Confined to oceanic regions, mainly tropical and subtropical montane but 2 spp. in
Greatest richness in W Pacific and temperate S Hemisphere. Mainly humicolous
Widespread in cool oceanic regions: W Europe, NW North America, NE Asia, S
Cool temperate to subarctic regions. Terricolous and strongly hygrophilous.
Arctic and alpine preferences: main diversity in the cool N Hemisphere, especially
Almost confined to montane forests around the W Pacific.
Plagiochila is most important genus with large numbers of species in tropical
montane rainforests. Very strongly represented in SE Asia and W Pacific.
Cosmopolitan, but greatest diversity in rainforest vegetation: SE Asia, S America.
Extremely diverse and important family, especially within the tropics; greatest
diversity in W Pacific, Indonesia and tropical America. In Europe almost confined
to the Atlantic seaboard. Mainly corticolous and epiphylious
Frullania, with over 400 species, is the most important genus. Greatest diversity
in the montane tropics of SE Asia and America. Mainly strongly photophilic
Comparatively few species, more or less evenly distributed. Soil-dwelling, mainly
Montane forests; SE Asia, S America, W Europe and N America. Hygrophilous,
Mediterranean type climate: W Mediterranean, S Africa, India. Xerophytic
tendencies. Includes some ‘weedy’ cosmopolitan species.
FAMILIES GENERA
Calobryales
2 2
Treubiales
1 2
Jungermanniales
39 families incl: The ‘leafy liverworts’.
Herbertaceae 4
W Europe.
Lepidoziaceae 24
and lignicolous.
Lophoziaceae 18
America, Australasia.
Jungermanniaceae 11
Gymnomitriaceae 3
W Europe.
Schistochilaceae 2
Lophocoleaceae 15 Cosmopolitan. Mainly ground-dwelling.
Plagiochilaceae 6
Radulaceae 1
Lejeuneaceae c. 70
Frullaniaceae 3
epiphytes but also epilithic etc.
Metzgeriales
5 families incl: Thalloid liverworts, strongly hygrophilous.
Dilaenaceae 11
riparian
Aneuraceae 2
mainly on soil, rocks and rotting wood
Metzgeriaceae 1 Widely distributed.
Marchantiales Thalloid liverworts
30
Anthocerotales Hornworts
5
ALGAE
Chlorophyta (Green Algae)
Widespread. Epiphylls maximum diversity in the W Pacific. Terrestrial
hygrophytes, epiphytes and epiphylls.
distributions. Many of the larger Chlorophyta are restricted
to either marine or freshwater conditions; a few are
sufficiently tolerant to be found in both environments. Table
7.5 lists the orders and constituent families of the larger
The class Chlorophyta is cosmopolitan in distribution and
Occurs in marine and brackish water, freshwater, and
terrestrial environments. It comprises approximately 1,040
species in 170 genera and contains eight orders (Silva,
1982) some of which have restricted geographical
57
Chlorophyta (excluding unicellular forms) and indicates
their broad geographical distributions and salinity
tolerances.
The largest family, the Cladophoraceae, occurs globally and
1. Biological Diversity
Table 7.4 Lichen diversity
REGION
USA and Canada
Australia
France
Sweden and Norway
West Indies
United Kingdom
New Zealand
India
Mexico
Philippines
Argentina
Sardinia
Hawaii
Tasmania
East Africa (macrolichens)
Central America
Guianas
New Guinea
Galapagos
Juan Fernandez
Ecuador
Table 7.5
Ulotrichales
Ulvales
Capsosiphonaceae
Percursariaceae
Ulvaceae
Monostromaceae
Prasiolales
Acrosiphoniales
Codiolaceae
Acrosiphoniaceae
Cladophorales
Cladophorales
Anadyomenaceae
Siphonocladales
Siphonocladaceae
Valoniaceae
Bryopsidales
Bryopsidaceae
Ostroebiaceae
Dichotomosiphonaceae
Caulerpaceae*
Udoteaceae
Codiaceae**
Dasycladaceae* **
GENERA
401
299
181
216
173
250
243
163
130
137
122
178
104
173
73
120
165
137
80
31
160
FAMILIES
GEN
11
SPP
50
in a wide range of salinities; the next largest family, the
Codiaceae, is restricted to the marine environment and does
not occur in the colder waters of the polar regions.
Temperature-dependent distribution is clearly seen at both
order and family levels. The Acrosiphoniaceae is restricted
to colder waters in contrast to the Siphonocladales,
Caulerpaceae and Udoteaceae’ which occur only in tropical
and subtropical waters.
Of the selected floras assessed for species diversity (Table
7.8), the North Atlantic, the tropical/subtropical western
Atlantic, and the Japanese region of the Pacific are the most
species-diverse. Although the green seaweed flora of
southern Australia is not so species-diverse, it probably
contains the highest number of endemics (46% of the total).
Particularly impoverished floras are those of the tropical
west coast of Africa (e.g. Gambia to Angola) and the west
coast of South America (e.g. Colombia, Peru), areas where
there are major cold water upwellings. Other impoverished
floras include those of small isolated islands (e.g.
Macquarie Island) and the polar regions.
Phaeophyceae (Brown Algae)
The Phaeophyceae are global in distribution, occurring in
polar, temperate and tropical zones. The brown algae are
principally marine plants, with only very few species in
freshwater. The class contains about 265 genera and in
excess of 1,500 species arranged in 14 orders (Wynne,
1982, Table 7.6).
Orders and families of larger green algae
DISTRIBUTION ECOLOGY
Global Mostly freshwater
Brackish water
Brackish water
Global Mostly seawater/brackish water
seawater/brackish water
Global Seawater/brackish water/freshwater
Seawater/brackish water
Seawater/brackish water
Seawater/brackish water
Cold waters
Cold waters
Cold waters
Global Seawater
Tropical/subtropical Seawater
Tropical Seawater
Tropical Seawater
Tropical Seawater
Global Seawater
Shells Seawater
Tropical/subtropical Seawater
Tropical/subtropical Seawater
Global Seawater
Tropical/subtropical Seawater
Notes: * Temperate in southern hemisphere, ** one relict genus in inland brackish water, *** not polar waters. GEN = genera, SPP = species.
58
Lower Plant Diversity
Table 7.6 Orders and families of brown algae
FAMILIES GENERA SPECIES DISTRIBUTION ECOLOGY
Ectocarpales 3 Global Marine
Ectocarpaceae 29 Global Marine
Ralfsiaceae 17 Marine
Sorocarpaceae 2 2 N Atlantic Marine
Chordariales 10
Myrionemataceae 11 Marine
Elachistaceae 5 Global Marine
Corynophloeaceae 5 Marine
Spermatochnaceae 5 Marine
Acrotrichaceae 1 1 N Atlantic Marine
Chordariaceae 29 Global Marine
Ischigiaceae 1 2 Limited Marine
Chordariopsidaceae 1 S Africa Marine
Notheiaceae 1 1 Australasia Marine
Splachnidiaceae 1 S Africa Marine
Cutleriales 1 3 Warm waters Marine
Tilopteridales 1 2 N Atlantic Marine
Sphacelariales 4 Global Marine
Sphacelariaceae 5 Global Marine
Stypocaulaceae 4 10 Global Marine
Cladostephaceae 1 N Atlantic/Australasia Marine
Choristocarpaceae 2 N Atlantic/Mediterranean Marine
Dictyotales 1 16 Tropical/subtropical * Marine
Sporochnales ‘1 Warm waters Marine
Desmarestiales 2 Cold waters Marine
Desmarestiaceae 3 Cold waters Marine
Arthrocladiaceae 1 N Atlantic/Mediterranean Marine
Dictyosiphonales ai Marine
Myriotrichaceae 1 N Atlantic/Mediterranean Marine
Giraudiaceae 1 1 N Atlantic/Mediterranean Marine
Striariaceae 9 Marine
Delameriaceae 4 Marine
Punctiariaceae 17 Temperate Marine
Chnoosporaceae 1 74 Tropical/subtropical Marine
Dictyosiphonaceae 2 N Hemisphere Marine
Scytosiphonales 1 8 Marine
Laminariales 4 Temperate/polar Marine
Chordaceae 1 2 N Atlantic Marine
Laminariaceae 15 Temperate/polar Marine
Lessoniaceae 8 NE Pacific/S Hemisphere Marine
Alariaceae 7 Temperate/polar Marine
Fucales 6 Marine
Fucaceae 7 N Hemisphere Marine
Brackish
Himanthaliaceae 1 1 NE Atlantic Marine
Hormoseiraceae 1 1 Australasia Marine
Phyllosporaceae 6 Australasia Marine
Sargassaceae 6 Tropical/temperate Marine
Cystoseiraceae 16 Tropical/temperate Marine
Durvilleales 1 1 4 Australasia/Antarctic Marine
Ascoseirales 1 1 1 Antarctic Marine
The more primitive orders (Ectocarpales, Chordariales) are
global in distribution, although some of the constituent
families, particularly the smaller ones (e.g., Sorocarpaceae)
are geographically restricted. The small orders Cutleriales,
Dictyotales and Tilopteridales are limited, respectively, to
59
warmer waters, the tropics and subtropics, and the North
Atlantic, while the Durvilleales and Ascoseirales occur only
in Australasia and Antarctica. The kelps (order
Laminariales) are disjunctly distributed in temperate waters
of both northern and southern hemispheres. In the most
1. Biological Diversity
Table 7.7
Bangiophycideae
Porphyridiales
Porphyridiaceae
Goniotrichaceae
Phragmonemataceae
Bangiales
Erythropeltidaceae
Bangiaceae
Boldiaceae
Compsopogonales
Rhodochaeteles
Florideophycideae
Nemaliales
Acrochaetiaceae
Batrachospermaceae
Lemaneaceae
Thoreaceae
Helminthocladiaceae
Nemaliaceae
Dermatonemaceae
Chaetangiaceae
Naccariaceae
Bonnemaisoniaceae
Gelidiaceae
Gelidiellaceae
Wurdemanniaceae
Cryptonemiales
Weeksiaceae
Dumontiaceae
Choreocolacaceae
Cryptonemiaceae
Corynomorphaceae
Pseudoanemoniaceae
Kallymeniaceae
Endocladiaceae
Crossocarpaceae
Gloiosiphoniaceae
Tichocarpceae
Pterocladiophilaceae
Peyssonneliaceae
Corallinales
Hildenbrandiales
Gigartinales
Gymnophleaceae
Gracilariaceae
Sebdeniaceae
Calosiphoniaceae
Petrocelidaceae
Phyllophoraceae
Gigartinaceae
Chondriellaceae
Polyideaceae
Nizymeniaceae
Rhizophyllidaceae
Acrotylaceae
Plocamiaceae
Phacelocarpaceae
Sarcodiaceae
Furcellariaceae
Solieriaceae
Hypneaceae
Rissoellaceae
Orders and families of red algae
FAMILIES GENERA SPECIES DISTRIBUTION
13
13
27
on
FH -NUOANN AWN
=
--NHANHON "ON DW
wo
- Oo
PH WV WANN OWNHNH HAHN NA
12
400
Global
Tropical/subtropical
Global
Global
Global
Global
S Hemisphere
Global
NW Pacific
Global
Juan Fernandez
S Australia
S Hemisphere
S Hemisphere
Mediterranean
60
ECOLOGY
Marine/freshwater
Marine/brackish/freshwater
Freshwater
Marine/freshwater
Marine
Marine/brackish/freshwater
Marine
Marine/brackish/freshwater
Marine/brackish/freshwater
Freshwater
Freshwater
Freshwater
Marine
Marine
Marine
Marine
Marine
Marine
Marine
Marine
Marine
Marine
Marine
Marine
Marine
Marine
Marine
Marine
Marine
Marine
Marine
Marine
Marine
Marine
Marine
Marine
Marine/brackish/freshwater
Marine
Marine
Marine
Marine
Marine
Marine
Marine
Marine
Marine
Marine
Marine
Marine
Marine
Marine
Marine
Marine
Marine
Marine
Marine
Marine
Lower Plant Diversity
Table 7.7 Orders and families of red algae (continued)
FAMILIES GENERA SPECIES DISTRIBUTION
Rhabdoniaceae 5
Cubiculosporaceae 1 1
Rhodophyllidaceae 3
Mychodeaceae 1 11
Dicranemaceae 4
Ahnfeltiales 1 1
Rhodymeniales 2
Rhodymeniaceae 30
Champiaceae 6
Palmariales 1 4
Ceramiales 4
Ceramiaceae 100
Delesseriaceae 100 300
Dasyaceae 12 100
Rhodomelaceae 100 500
S Australia
S Australia
Global
Global
Global
ECOLOGY
Marine
Marine
Marine
Marine
Marine
Marine
Marine
Marine
Marine
Marine
Marine
Marine/brackish* /freshwater*
Marine/freshwater*
Marine
Marine/brackish*
Note: * A few species only.
advanced order, the Fucales, the family Cystoseiraceae
occurs widely in the tropical and temperate zones, whereas
the family Sargassaceae is mostly confined to the tropical
waters. Other families have more circumscribed
distributions, the Hormoseiraceae and Phyllosporaceae
occurring only in Australasia; the Fucaceae is restricted to
the northern hemisphere and the monotypic Himanthaliaceae
is endemic to the north-eastern Atlantic.
Brown algae attain greatest species richness in the Japanese
region of the Pacific, the North Atlantic and, to a lesser
extent, southern Australia. The last region, however, is
probably highest in endemics, with 18% of generaand 54%
of species endemic while only 26% of the flora comprises
widespread species. Species-depauperate floras are, as in
the Chlorophyta, those in cold-water upwelling areas (e.g.
Angola, Colombia and Peru), on isolated islands
(Ascension, St Helens), or a combination of both (e.g.
Macquarie Island).
Rhodophyta (Red Algae)
The Rhodophyta is the largest of the three main seaweed
groups, with over 555 genera (Dixon, 1982, Table 7.7); it
contains more species than the Chlorophyta (Green) and
Phaeophyceae (Brown) together. The Rhodophyta is divided
into two subclasses, the subclass Bangiophycideae, the
smaller of the two, occurs throughout the world in marine,
brackish and freshwater environments. The subclass
Florideophycideae comprises eight orders and is
predominantly marine. None of the orders is clearly
circumscribed geographically; some small families (e.g.
Mychodeaceae, Phacelocarpaceae) are restricted in
occurrence to Australia or the southern hemisphere
generally. The larger families are widely distributed.
As with the Green and Brown Algae, the most species-rich
floras are those of the Japanese Pacific region, the tropical
and subtropical western Atlantic, and the North Atlantic
(including temperate and arctic regions). Other rich floras
61
of Red Algae are those of California and Chile. Although
precise data are not available for southern Australia, it is
probably also species-rich, with 75 % of species and 30% of
genera endemic to the area. Species-poor floras are those
referred to previously on the tropical west coasts of Africa
and South America where there are cold-water upwellings.
Red algae floras decrease in species abundance in cool
waters.
General remarks on marine algal floras
The most species-rich algal flora assessed is that of the
Japanese region of the Pacific (1,503 species, Table 7.8).
The North Atlantic and tropical and subtropical western
Atlantic are also species-rich, with over 1,000 species
recorded.
In the North Atlantic, eastern and western seaboards differ
in diversity. The western (American) coastline is relatively
species-poor; 65% of the North Atlantic flora is restricted
to the eastern (European) coast, 35% is common to both
coasts, and only 5% restricted to the American coast. A
reduction in species also occurs from south to north, with
the Arctic flora the least diverse and characterised by
hardy, cosmopolitan species and very low endemism. The
flora of the British Isles is relatively species-rich (over 700)
exceeding that north western Pacific America and one of the
richest of the 18 floras assessed. The flora of the Eastern
Mediterraneanis probably much richer than indicated by the
430 species listed for Aegean Greece. The latter flora is
fairly high in endemics (20% of the total species, while
28% of all species are Méediterranean-Atlantic in
distribution).
The flora of the tropical and subtropical regions of the
Atlantic contrast with those in higher latitudes to the north
in having the most species-diverse area on the western
(American) side, where 1,058 species are recorded. On the
eastern (tropical African) side the number is only about
300. Similar comparison of the Chlorophyta (green algae)
1. Biological Diversity
and Phaeophyta (brown algae) floras (groups having better
data for subtropical and tropical Africa) shows 253 species
of green algae from the west and 153 from the east, and
150 species of brown algae from the west and 125 from the
east. The flora of tropical west Africa contains 56% of
species that also occur in the Indian Ocean and 58% of
species in the Pacific Ocean.
Moderately rich floras are those of Chile (temperate),
North-west America (temperate), California (subtropical)
and tropical East Africa. The flora of southern Australia
probably also falls into this group, but is probably much
higher in endemics.
Species-poor floras generally occur in polar waters, and on
isolated islands - the further from the nearest landmass the
poorer the flora (e.g. St Helena with only 68 species).
Potential for endemism exists in water masses isolated from
the main oceans, such as the Mediterranean Sea (which only
has very small water exchange with the Atlantic Ocean), the
Black Sea (for similar reasons), and the Caspian Sea (now
completely isolated and brackish, but retaining an
Table 7.8 Diversity of marine algal (seaweed) floras
CHLOROPHYTA PHAEOPHYCEAE RHODOPHYTA TOTAL
FLORA GENERA SPECIES GENERA SPECIES GENERA SPECIES GENERA SPECIES
Japan 60 234 108 379 267 900 475 1503
N Atlantic 67 253 127 324 193 539 387 1116
W Atlantic 64 253 63 150 134 655 321 1058
Chile 31 131 60 140 150 480 241 751
California 22 72 69 137 186 459 279 668
E Africa 30 159 29 118 121 366 180 643
NW America 51 117 66 143 161 373 273 635
Antarctica 88 118 357 563
S Africa 547
E Mediterranean 30 73 49 90 131 267 210 430
Viet Nam 115 86 223 424
Red Sea 30 92 34 118 79 173 143 383
Tropical W Africa 19 59 22 42 88 198 129 299
Angola 8 34 18 22 71 140 97 196
Peru 20 20 107 156
Colombia 12 23 13 21 46 79 71 123
Macquarie | 12 15 25 28 46 60 81 103
St Helena 10 13 9 10 34 45 53 68
Ascension | 9 14 11 15 16 23 35 52
S Australia 39 119 104 231
Tropical/subtropical
Charophyta (Stoneworts)
The charophytes or stoneworts are a very distinctive group
of macrophytic green algae that occur from Spitzbergen in
the north (80°N) to the Kerguelen Islands in the south (c.
50°S). A few are restricted to brackish water but the large
majority are widely distributed in such freshwater habitats
as ponds, lakes, ditches, temporary pools, streams, rivers
and swamps. The six extant genera are placed in two tribes:
the Chareae - Chara, Lamprothamnium, Nitellopsis and
Lychnothamnus; and the Nitelleae - Nitella and Tolypella.
It is difficult to undertake a biogeographic analysis based on
charophyte species because of the current uncertainty
surrounding taxonomic limits and the ranking of
62
impoverished seaweed flora).
An important characteristic of many tropical and subtropical
regions is the occurrence of coral reefs; algae are a major
constituent of these long-stable ecosystems and the sheltered
lagoons they protect. Coral reefs support a unique and
generally diverse algal flora that includes many crustose
coralline algae whose numbers are likely to increase with
further study. Mangrove areas are also restricted to the
tropics and subtropics and support a well-defined and
interesting algal vegetation, contrasting with that of
saltmarshes in the temperate zones, which are generally
more species-poor. Sandy coastlines are floristically
depauperate areas and often form barriers to seaweed
dispersal. Some anthropomorphic changes to the coastline
involving creation of additional habitats have locally
enhanced species diversity; pollution, in contrast, has
reduced species diversity, especially in lagoons, mangrove
areas and coral reefs. In the latter, pollution-tolerant weedy
species appear to outcompete and replace pollution-sensitive
species. Land reclamation, rice-paddies and salt-pan
development have led to the loss of algal habitat in many
coastal areas in the tropics.
infrageneric taxa. In Wood’s world monograph on the
group (Wood, 1965) the infrageneric taxa were divided into
sections, species, subspecies, varieties and forms. He
considered morphologically similar monoecious and
dioecious taxa to be ‘species pairs’ and combined them.
Other charologists do not accept Wood’s views on merging
monoecious and dioecious taxa and continue to regard them
as distinct.
Unlike most other groups of freshwater algae sufficient
regional information exists on the distribution of
charophytes to allow for global analysis. In carrying out
such an analysis Khan and Sarma (1985) used Wood’s
classification but did not recognise the merging of
monoecious and dioecious taxa. They included charophytes
described after 1965 and taxa reduced by Wood to
synonymy but subsequently shown to be distinct. Khan and
Sarma recognised 440 taxa, of which 274 were known from
only one region or continent (‘endemics’). For assessing the
geographical distribution of taxa eight broad zones
(regions/continents) were recognised: North America, South
America, Africa, Europe, Asia (including Japan but
excluding India), India, Pacific Island region, and Australia.
Antarctica was not included as it is the only continent for
which charophytes have yet to be reported.
Table 7.9 Stonewort diversity
REGION GENERA SPECIES ENDEMIC
SPECIES
North America 4 114 50
Asia 5 122 48
Africa 4 116 42
Europe 6 91 41
Australia 5 62 25
South America 5 89 25
India 6 125 23
Pacific Region 4 72 19
World 6 440 274 *
Source: Khan, M. and Sarma, Y.S.R.K. 1984. Cytogeography and
Cytosystematics of Charophyta. In: Irvine, D.E.G. and John, D.M.
(Eds), Systematics of the Green Algae. Academic Press, London and
Orlando.
Note: * Majority of the remainder (c. 166) have a restricted
distribution (normally two or three regions/continents) and about seven
are to be regarded as cosmopolitan. Fewer than a dozen taxa have been
published since 1985 and most are from underworked regions (e.g.
South America; Asia, especially China).
It is impossible to determine to what extent tabulated
estimates are significant or simply reflect collecting. Europe
is one of the most intensively collected regions and so the
lower numbers reported are likely to represent a real
difference in diversity. The general unsatisfactory state of
the taxonomy will continue to hamper biogeographical
analysis.
Charophytes form extensive and sometimes diverse
associations in marl rich water bodies and are especially
sensitive to nutrient enrichment or eutrophication. In some
countries they have become dramatically less common and
more restricted in distribution as a result of nutrient
enrichment primarily from agricultural sources. This would
seem to be the main threat to these algae along with the
general loss of aquatic habitats through land reclamation.
Brackish-water lagoons is an example of a habitat under
threat in many countries and one the genus Lamprothamnion
is almost wholly confined to it. In the British Isles this is
the only charophyte protected by government legislation
63
Lower Plant Diversity
although several freshwater species may also be under
threat and have been recommended for protection. The
conservation status of charophytes is difficult to determine
without considerably more information on habitat
requirements.
Other groups of algae
Comments on the diversity and global distribution of most
groups of microalgae are not possible because of inadequate
knowledge of the algal floras of the world. A reasonable
coverage exists for a few regions but only for fairly well-
defined algal groups such as the desmids (Division
Chlorophyta, Order Desmidiales) and the diatoms (Division
Bacillariophyta). Only a few attempts to analyse and
interpret regional distribution patterns go so far as to
consider the wider distribution of individual taxa. Doubt is
often attached to the reliability of published lists so that the
findings of regional comparisons need to be treated with
caution. Frequently, ‘regional endemics’ have had to be
reduced to synonymy because the describing authors failed
to take adequate account of the taxonomic literature
covering other regions. Sometimes the converse is true, and
endemics are not recognised because they are incorrectly
attributed to an extant taxon using identification guides
written for another region. If progress is to be made it is
essential to have sounder species concepts, more accurate
identification, and considerably more information on the
algal floras of under-collected parts of the world.
References
Dixon, P.S. 1982. Rhodophycota. In: Parker, S.P. (Ed.),
Classification of Living Organisms. McGraw Hill, New York.
Pp.62-79.
Galloway, D.J. 1992. A lichenological perspective. Biodiversity and
Conservation: submitted September 1991.
Hawskworth, D.L. and Ahti, T. 1990. A bibliographic guide to the
lichen floras of the world, 2nd edn. Lichenologist 22:1-78.
John, D.M. 1986. The algal flora: its analysis and biogeography. In:
John, D.M., The Inland Waters of Tropical West Africa. E.
Schweizerbart’sche, Stuttgart. Pp.133-160.
Khan, M. and Sarma, Y.S.R.K. 1984. Cytogeography and
Cytosystematics of Charophyta. In: Irvine, D.E.G. and John, D.M.
(Eds), Systematics of the Green Algae. Academic Press, London
and Orlando. Pp.303-330.
Silva P.C. 1982. Chlorophycota. In: Parker, S.P. (Ed.), Classification
of Living Organisms. McGraw Hill, New York. Pp.133-161.
Wood, R.D. 1965. In: Wood, R.D. and Imahori, K. (Eds), A Revision
of the Characeae, Part I. Cramer, Weinheim.
Wynne, M.J. 1982. Phaeophyceae. In: Parker, S.P. (Ed.),
Classification of Living Organisms. McGraw Hill, New York.
Pp.115-125.
Chapter abridged from material contributed by the following
staff of the Department of Botany, The Natural History
Museum. (London):
Alan Eddy (Bryophytes); D.J. Galloway (Lichens); David
M. John (Algae); lan Tittley (Green Algae).
1. Biological Diversity
8. HIGHER PLANT DIVERSITY
The higher plants, characterised by vascular tissue and
reproducing either by spores, cones, or flowers, dominate
the world’s flora and vegetation. Along with the bryophytes
(Chapter 7), they develop from an embryo resulting from
the sexual fusion of cells. They consist of three groups:
e@ the pteridophytes or ferns and fern allies, such as
clubmosses, horsetails, quillworts and whiskferns
e the gymnosperms, mainly the conifers and cycads
e the angiosperms or flowering plants.
THE GROUPS OF HIGHER PLANTS
Pteridophytes
Estimates of the total number of ferns and their allies vary
between 10,000 and 13,000 species but is probably close to
12,000, the majority of which are native to the moist
tropics.
The so-called ‘fern allies’ probably do not form a natural
group but rather represent the end points of several distinct
evolutionary lineages. Like the true ferns, they reproduce
by spores. The earliest known vascular land plants belong
to this group. These psilophytes (Psilophyta), which
dominated the landscape during the Silurian and Devonian
around 400 million years ago (Mya), are all but extinct;
they are only represented by two relict genera - Psilotum
(tropics) and Tmesipterus (Australia, New Zealand, South
Pacific). Psilotum is extremely primitive, lacking both roots
and leaves.
Today, the lycopods (Lycopodiophyta) are represented by
only five relict genera (Isoetes, Lycopodium, Phylloglossum,
Selaginella, and Stilites), but their fossil record extends
back to the Carboniferous (c. 300 Mya), when they formed
the dominant vegetation. These extinct forms grew to 40m
high and had a stem diameter of 2m; their remains form
part of the coal reserves we rely on today.
The horsetails and scouring rushes (Sphenophyta) are
another ancient group, and are also all but extinct. They are
represented by a single genus, Equisetum, containing some
15 species found throughout the world, but especially well
represented in North temperate bogs.
The true ferns (Pteridophyta or Filicophyta) are much more
diverse than are the fern allies. They show great range of
form, from the tiny, delicate filmy ferns
(Hymenophyllaceae) to tropical tree-ferns (Cyatheaceae and
Dicksoniaceae) more than 15m tall; leaves vary in length
from Smm to 10m. Ferns are cosmopolitan in distribution
but are scarce in arid zones and occur in greatest numbers
in the moist tropics, where they often grow epiphytically.
It has been estimated that 12.5% of the world’s fern species
are to be found in Papua New Guinea (Johns and Bellamy,
1979), and 10% in India (Dixit 1984). Some species have
a very wide distribution, notably Bracken Pteridium
aquilinum, which is found throughout the temperate zones
and over much of the tropics, while other species are
extremely limited in their distribution.
64
Gymnosperms
The gymnosperms are trees (or occasionally shrubs) whose
seeds lack the covering characteristic of the flowering
plants. They include some 500 species of conifer, 100
species of cycad, and a few other small but scientifically
fascinating families. They first appear in the fossil record
in the Carboniferous (c. 300 Mya) as the so-called ‘seed
ferns’ (which were not true ferns at all, but intermediates
between ferns and gymnosperms). Gymnosperms dominated
the earth until the rise of the flowering plants.
Conifers occur worldwide, but they reach their greatest
diversity of species and genera in parts of Oceania and on
the margins of the Pacific Ocean. They are the softwoods
of commerce and are widely grown for timber and
ornament. A conifer from the western USA, the Giant
Sequoia Sequoia sempervirens is the tallest tree in the
world, reaching a height of 110m; another conifer from
western USA, the Bristlecone Pine Pinus aristata is thought
to include the oldest living individual trees on earth, some
being 4,900 years of age. The largest genera are the pines
Pinus, firs Abies, and spruces Picea, which form extensive,
economically important forests in the boreal zone of Eurasia
and North America and in the mountains of the northern
hemisphere. The podocarps Podocarpus are widespread in
tropical and subtropical forests of the southern hemisphere.
Locally, other genera are prominent, such as kauri pines
Agathis (exploited for resin) in wet forests from Malesia to
New Zealand, and Chinese Fir Cunninghamia lanceolata,
the major timber tree of South and West China.
Cycads, palm-like tropical trees, occur mostly in Central
and South America, South Africa, and from Southeast Asia
to Australasia. They include the Sago-palms Cycas, an
ancient group which originated at least 240 Mya and are
thus of considerable scientific interest. Many of them are
highly restricted in their distribution and are of great
conservation concern.
Other gymnosperms include the famous maidenhair tree
Ginkgo biloba, an isolated, ancient relict species native to
China, the yews Taxus (source of the promising drug taxol)
and their allies; joint-pines Ephedra, leafless ‘switch plants’
of scrub and semi-desert, Gnetum, mostly lianes of moist
tropical forests, and the remarkable Welwitschia bainesii,
which looks like a great woody turnip bearing only two
huge, strap-shaped leaves and a cluster of either male or
female cones, restricted to the coastal fog-belt of the Namib
desert of Angola and Namibia. As a general rule, however,
Africa has a very poor gymnosperm flora.
Angiosperms
The flowering plants, or Angiosperms, are an extremely
diverse group of plants, containing some 250,000 species
(see Table 8.2). From their first appearance in the fossil
record around 135 million years ago, they evolved quickly
and have come to dominate all other land plants, except in
certain habitats (such as the boreal region, in which
gymnosperms dominate). Most of our food comes from
angiosperms, as do many spices, drugs, poisons, fibres,
building materials. Many angiosperms are much utilised for
their valuable timber (see Part 2).
Angiosperms are seed-producing plants that bear flowers
that are often insect- or bird-pollinated. The plants range in
size from 1mm (Wolffia spp.) to over 100m tall (Eucalyptus
regans from Tasmania). The flowers can reach over 1m
across (Rafjlesia arnoldii from Sumatra and Borneo).
Estimates of the number of flowering plant species vary
between 240,000 and 750,000, but most botanists accept
250,000 species as the best figure. These species are
grouped into some 17,000 genera. Despite an enormous
diversity of growth form and floral structure, the number of
flowering plant families recognised is relatively small. It
has varied over the years from 200 to over 600, but there
is now general agreement on a basic 300-400 ‘core’ families
of flowering plants. Many of these families, such as
Compositae (daisy and dandelion family) and Cruciferae
(cabbage family) are natural units, and can be recognised
without too much difficulty by the non-botanist, while
others are characterised by more technical features not
easily discernible by the layman.
Families vary greatly in the number of species they contain:
on the one hand there are massive families like Orchidaceae
(orchid family) with 25,000-35,000 species and
Leguminosae (pea and bean family) with about 14,500
species (see Table 8.2). In fact, only 31 families contain
62% of known flowering plant species. At the other
extreme are the 36 families with a single species, such as
the Adoxaceae, the family of the well known North
European woodland flower, Moschatel Adoxa
moschatellina.
The grouping of these families into higher taxonomic levels
such as orders and subclasses is somewhat more
problematical, reflecting uncertainty about the fundamental
evolutionary relationships between families. A commonly
used scheme (after Cronquist, 1981) is presented in Table
8.2.
THE DISTRIBUTION OF HIGHER PLANTS
Higher plants occur in virtually all ecosystems of the world,
even in the sea, but their distribution is very uneven. Two-
thirds of the world’s flowering plants are tropical,
emphasising the great importance of plant conservation in
the tropics. Many large or economically important families
such as Annonaceae (custard-apple family), Lauraceae
(cinnamon family), Moraceae (fig family), Dipterocarpaceae
(dipterocarp family), Ebenaceae (ebony family) and
Meliaceae (mahogany family) are almost entirely restricted
to the tropics. This contrasts with the distribution of those
who study plants, for specialists in plant taxonomy work
mostly in Europe or the USA. The richest continent for
plants, and still the least explored botanically, is South
America, home to perhaps as much as one-third of the
world’s higher plants.
Table 8.1 gives an assessment of the numbers of species of
higher plants in various regions of the world. Some of the
figures, however, are provisional estimates that need to be
65
Higher Plants
treated with caution. It must be emphasised also that the
species concept used varies from one region to another,
which means that any comparison of the numbers of plants
between regions must be done with care.
In particular, the differences in species richness between the
regions of the world shown in Table 8.1 may be somewhat
exaggerated. The species concept commonly used in Latin
America, for example, tends to recognise more species,
based on characters visible in the field, than the taxonomy
of botanists working on the Malesian region. South America
is still the continent with the most plants, but the
differences between this region and tropical Asia or Africa
may in time be found to be less than suggested. For
example, estimates of the size of the flora of Colombia, a
territory with high levels of species diversity and
endemism, fell over a ten-year period from 45,000 (Prance,
1977) to 35,000 (Forero 1988).
A degree of convergence is apparent. In 1985, IUCN cited
figures of 20,000 species in North America and 11,300 in
Europe (Davis ef al., 1986). In Table 8.1 the estimate for
North America has dropped to 17,000, following revised
estimates by the Flora of North America workers, while
that for Europe has risen to 12,500, following predictions
based on the many species added to the recently revised
first volume of Flora Europaea. It is fair to assume that
North America does have more plants than Europe, but
further convergence between the two figures is likely.
These changes in numbers of species do not result strictly
from extinctions or the evolution of new species, although
both of these processes are happening. In most cases, they
result from decisions of botanists as to the delimitation of
individual species. Many species in a flora are not clearly
defined entities, as is, for example, the Gingko tree Ginkgo
biloba, but are members of a complex group of species
between which differences may be small. This is
particularly true of some tropical and Mediterranean floras,
where many species are extremely difficult to identify in the
field. At the same time, collaboration between botanists
who study the floras of different continents (facilitated by
modern information technology and electronic data retrieval
systems) is helping to rationalise and standardise the
classification of plants that have in the past been treated as
distinct species in different regions. Opinions will naturally
vary as to the use of the rank of species, subspecies or
merely variety.
Individual botanists tend to study either the plants of a
particular country or the members of a particular family.
Consequently, few data are available as to the numbers of
species in individual habitats. Nevertheless, some general
points can be made. Tropical forests, especially moist
forests, are of enormous importance as habitats for plants.
The species diversity of these forests, alongside fossil
evidence, has led many botanists to argue that the flowering
plants evolved in tropical forests, although it is more likely
that they represent a ‘museum’ of evolution (Stebbins,
1974). Probably half or slightly under half of all higher
plant species are restricted in the wild to tropical forests, a
proportion that may be a little lower than that of animals
because of the exceptional plant richness of Mediterranean
ecosystems, a richness that is not reflected in faunal
1. Biological Diversity
Table 8.1 Distribution of higher plants
by continents
Latin America (Mexico through S America) 85,000 '
Tropical & Subtropical Africa 40,000 - 45,000
North Africa 10,000 ?
Tropical Africa 21,000 *
Southern Africa 21,000 *
Tropical & Subtropical Asia 50,000 ®
India 15,000 °
Malesia 30,000 ’”
China 30,000 ®
Australia 15,000 °®
Caribbean
Pacific
North America 17,000 ®
Europe 12,500 '°
Sources: ' Gentry, A.H. 1982. Neotropical floristic diversity:
phytogeographical connections between Central and South America,
Pleistocene climatic fluctuations, or an accident of the Andean
orogeny? Annals of the Missouri Botanical Garden 69:557-593 .* Based
on figures for the size of country floras given in Quezel, P. 1985.
Definition of the Mediterranean region and the origin of its flora. In:
Go6mez-Campo, C. (Ed.), Plant Conservation in the Mediterranean
Area. Junk. P.17. > Estimate by A.L. Stork, quoted by Peter Raven,
pers. comm., 1991. * Cowling, R.M. et al. 1989. Patterns of plant
species diversity in southern Africa. In: Huntley, B.J. (Ed.), Biotic
Diversity in Southern Africa: concepts and conservation. Oxford, Cape
Town. * From Raven, P.H. 1987. The scope of the plant conservation
problem worldwide. In: Bramwell, D. et al. (Eds), Botanic Gardens
and the World Conservation Strategy. Academic Press. Pp.19-29. ©
From Davis, S. et al. 1986. Plants in Danger: What do we know?
TUCN, Cambridge and Switzerland.’ M.M.J. van Balgooy, Leiden, in
lit. to J.R. Akeroyd, August 1991. * Prof. Wang Siyu, Beijing, in lin.
to J.R. Akeroyd, October 1991. Nancy Morin, pers. comm. via Peter
Raven, 1991. '° Estimate by J.R. Akeroyd, based on Flora Europaea,
1964-80, and the revision of Volume 1, in press.
Note: ‘Malesia’ consists of the nations of Malaysia, Brunei, Indonesia,
Philippines and Papua New Guinea.
diversity. It is estimated that the Mediterranean basin has a
flora of 25,000 species of higher plants (Quezel, 1985), a
high proportion of which are endemic. The other regions of
the world with a Mediterranean climate - the Cape Province
of South Africa, SW Australia, California, and Central
Chile - are also rich in endemics.
Patterns of plant distribution
Typical of most, but not all, groups of organisms, the
diversity of higher plants increases as one moves from the
poles to the equator. Plant species diversity, however,
varies markedly on smaller scales. Between 40 and 100 tree
species may occur on one hectare of tropical moist forest in
Latin America, compared to 10-30 per hectare in forests in
eastern North America. In a study done near Iquitos, Peru,
Gentry found approximately 300 tree species per hectare
with trunks greater than 10cm in diameter (Gentry, 1988).
Myers (1990) has estimated that 18 places on earth (termed
‘Hot-Spots’) support nearly 50,000 endemic plant species -
about 20% of the world’s total flora - but comprise only
0.5% of the earth’s surface. These 18 places, which range
widely in scale, are as follows: Atlantic coast of Brazil,
California Floristic Province, Cape Floristic Province,
Central Chile, Colombian Choco, Eastern Arc forests of
Tanzania, Eastern Himalayas, Céte d’Ivoire, Madagascar,
New Caledonia, Northern Borneo, Peninsular Malaysia,
Philippines, South Western Australia, Sri Lanka, Western
Amazonia uplands, Western Ecuador, and the Western
Ghats. This and other approaches to distinguishing areas of
high diversity are discussed further in Chapter 15.
Although the hot-spots sensu Myers are not defined by
habitat, they can be considered in such terms. Six units -
the Atlantic coast of Brazil, the Colombian Choco,
Northern Borneo, Peninsular Malaysia, the Philippines and
the Western Amazonia uplands - are areas of which the
natural vegetation cover. (now severely degraded) is almost
entirely tropical rain forest, a large proportion of it lowland
forest. Two more units - the Eastern Arc forests of
Tanzania and the Western Ghats in India - represent areas
of tropical montane forest. The vegetation of Western
Ecuador is essentially a mixture of both (Gentry, 1991).
Madagascar, Cote d’Ivoire and Sri Lanka each have a range
of habitats but those with by far the richest floras are the
tropical moist forests. The Eastern Himalayas are a region
of subtropical to warm-temperate forests, and New
Caledonia has a wide range of tropical habitats
(Schneckenburger, 1991). The four other units - the
California and Cape Floristic Provinces, Central Chile and
SW Australia -are regions of predominantly Mediterranean
vegetation.
Geopolitical distribution of plant diversity
Table 8.3 is a new compilation of higher plant richness and
endemism assessed on a territorial basis. The associated
figures are based on selected data from this table, and
illustrate the approximate percentage of country floras
composed of single-country endemic species (Fig. 8.1) and
the relative species richness of different countries. The 25
most species-rich countries are represented in Fig. 8.2 and
countries grouped by continent in Figs. 8.3-8.8 (note that
graph scales differ between continents).
It should be noted that these data reflect the size and
topographic complexity of the countries represented, in
addition to diversity per unit area as a function of climatic
and other factors. Nevertheless, the figures do confirm the
great floristic richness of the regions of moist tropical
forest. Territories that lie along the equatorial zone of moist
trade winds can have enormous numbers of species,
especially in South America: Venezuela has 15,000-25,000,
Colombia has 35,000, Brazil may have as many as 55,000
flowering plant species. African countries show a similar
high level of diversity, although numbers of species are not
as great as in South America, perhaps because of
prehistoric climatic fluctuation. Cameroon has an estimated
8,000 flowering plant species, Gabon 6,000-7,000 and
Tanzania 10,000. Floras in SW Asia are intermediate in
size between those of Africa and South America: there are
an estimated 20,000 flowering plant species in Indonesia
and 12,000 in both Malaysia and Thailand.
Amongst the richest floras are those of larger oceanic
islands in tropical and warm-temperate latitudes. Cuba has
a flora of 6,499 higher plant species, 3,233 of them
endemic; Japan has 5,372 species, some 2,000 of them
endemic; New Caledonia has 3,094 species, 2,480 of them
endemic; New Zealand has 2,371 species, 1,942 of them
endemic. The richest island flora is probably that of
Madagascar, estimated at up to 10,000 species, with
perhaps as many as 8,000 endemics. These include eight
endemic families of flowering plants, most notably the
spiny, rather cactus-like Didiereaceae that are a major
constituent of the vegetation in the drier parts of the island.
Smaller oceanic islands, even in the tropics, have small
floras due to the problems of long-distance dispersal for
plants, but the low total number of species frequently
includes a large endemic element. Mauritius, including
Réunion, has a native flora of 878 higher plant species, of
which 329 are endemic; Socotra has 788 flowering plants,
268 of which are endemic; St Helena has a native flora of
just 89 species, but 74 of these are endemic. Even some of
the very tiny atoll territories in Oceania usually have one or
a few endemic higher plants.
Drier tropical and subtropical regions, on the other hand,
have relatively poor levels of floral diversity when assessed
purely on a numerical basis. Most of the arid sub-Saharan
territories of the Sahel belt have smaller floras than have
many countries in N. Europe: for example, Burkino Faso
(1,100 higher plant species), Chad (1,600 species), Mali
(1,741 species) and Niger (1,178 species). These territories
have but a tiny number of endemics, perhaps no more than
a dozen between them. That is not to say that the Sahel
flora is not important, for it contains potentially valuable
drought-resistant and economic plants. They certainly show
a good deal less floristic diversity than the territories of the
Mediterranean region (noted above). Several of the
territories that border its shores have very high floral
diversity: Greece has 4,900 flowering plants, 742 of them
endemic; Spain about the same number, 941 of them
endemic; and Turkey 8,472 with 2,651 endemics. These
figure compare favourably with those from many tropical
territories, although they also reflect more thorough levels
of floristic exploration.
67
Higher Plants
References
Airy Shaw, H.K. (ed.). A Dictionary of the Flowering
Plants and Ferns. Eighth Edition. Cambridge Univ.
Press. 1245 pp.
Cowling, R.M. et al. 1989. Patterns of plant species
diversity in southern Africa. In: Huntley, B.J. (Ed.),
Biotic Diversity in Southern Africa: concepts and
conservation. Oxford, Cape Town.
Cronquist, A. 1981. An Integrated System of Classification of
Flowering Plants. Columbia University Press, NY.
Davis, S. et al. 1986. Plants in Danger: What do we know? TUCN,
Cambridge and Switzerland.
Dixit, R.D. 1984. A Census of the Indian Pteridophytes. Botanical
Survey of India, New Delhi.
Forero, E. 1988. Botanical exploration and phytogeography of
Colombia: past, present and future. Taxon 37:561-566.
Gentry, A.H. 1982. Neotropical floristic diversity: phytogeographical
connections between Central and South America, Pleistocene
climatic fluctuation, or an accident of the Andean orogeny? Annals
of the Missouri Botanical Garden 69:557-593.
Gentry, A.H. 1988. Tree species richness of upper Amazonian forests.
Proceedings of the National Academy of Sciences 85:156-159.
Gentry, A.H. 1991. Biological extinction in western Ecuador. Annals
of the Missouri Botanical Garden 78:273-295.
Johns, R.J. and Bellamy, A. 1979. The Ferns and Fern Allies of Papua
New Guinea. Papua New Guinea Forestry College.
Myers, N. 1990. The biodiversity challenge: expanded Hot-Spots
analysis. The Environmentalist 10(4):243-255.
Prance, G.T. 1977. Floristic inventory of the tropics: where do we
stand? Annals of the Missouri Botanical Garden 64:659-684.
Quezel, P. 1985. Definition of the Mediterranean region and the origin
of its flora. In: Gmez-Campo, C. (Ed.), Plant Conservation in the
Mediterranean Area. Junk. P.17.
Raven, P.H. 1987. The scope of the plant conservation problem
worldwide. In: Bramwell, D. et al. (Eds), Botanic Gardens and the
World Conservation Strategy. Academic Press. Pp.19-29.
Schneckenburger, S. 1991. Neukaledonien. Pflanzenwelt einer
Pazifikinsel. Palmengarten Sonderheft 16. Palmengarten, Frankfurt.
Stebbins, G.L. 1974. Flowering Plants. Evolution above the species
level. Edward Arnold. Pp.165-170.
Based on a document written by John Akeroyd and Hugh
Synge.
1. Biological Diversity
Table 8.2 Vascular plants: a summary of systematic diversity
MAJOR GROUP (CLASS)
SUBCLASS
ORDER
FAMILY GENERA SPECIES DISTRIBUTION
Pteridophytes \
Lycopodiaceae 4 587 cosmopolitan
mainly tropical, with some temperate
Selaginellaceae 1 725 species
lsoetaceae 1-2 77-80 temperate and tropical (aquatic)
Equisetaceae 1 22 cosmopolitan, except Australasia
Psilotaceae 2 3-10 tropical and subtropical
True ferns
Ophioglossaceae 3 81 temperate with some tropical
mostly Old World tropical; some New World
Marattiaceae 4 204 tropical
Osmundaceae 3 18 temperate and tropical
mostly Old World tropical; some New World
Plagiogyriaceae 1 36 tropical
Schizaeaceae 5 143 pantropical
Adiantaceae 38 Tes pantropical; subtropical; warm temperate
Parkeriaceae 1 4 pantropical
Vittariaceae 9 113 pantropical
Pteridaceae 7 259 pantropical
Marsileaceae 3 67 temperate and tropical
Hymenophyllaceae 5 600 pantropical
Hymenophyllopsidaceae 1 8 northern South America
Stromatopteridaceae 1 1 New Caledonia
Matoniaceae 2 4 Malesia
Gleicheniaceae 2 140 pantropical
Cheiropleuriaceae 1 1 tropical Asia and Malesia
Dipteridaceae 1 8 tropical Asia; Malesia; Australia; Fiji
Polypodiaceae 40 1,068 pantropical; subtropical; some temperate
Metaxyaceae 1 1 pantropical
Loxsomataceae 2 4 New World tropical; New Zealand
Thyrsopteridaceae Z 6 pantropical
Dicksoniaceae 3 41 pantropical
Lophosoriaceae 1 1 New World tropical
Cyatheaceae 4 623 pantropical
pantropical; some in subtropical and
Thelypteridaceae 30 1,000 temperate
Dennstaedtiaceae 18 486 pantropical
Aspleniaceae 14 711 pantropical; subtropical; some temperate
Woodsiaceae 18 705 pantropical; some temperate
Tectariaceae 19 431 pantropical
Dryopteridaceae 20 464 temperate; tropical
Lomariopsidaceae 8 615 pantropical
Davalliaceae 6 218 pantropical
Blechnaceae 8 238 pantropical
Salviniaceae 1 10 pantropical; subtropical; a few temperate
Azollaceae 1 6 pantropical; subtropical; some temperate
Gymnosperms - Cycads
Zamiaceae 8 80 tropical and subtropical
Madagascar; eastern and Southeast Asia;
Cycadaceae 1 20 Indomalaysia; Australia; Polynesia
Stangeriaceae 1 1 South Africa
Boweniaceae
Gymnosperms - Conifers
Northern Hemisphere, south to Sumatra,
Pinaceae 10 250 Java, Central America and West Indies
Northern Hemisphere, south to Celebes and
Taxaceae 5 20 Mexico; one species in New Caledonia
Taxodiaceae 10 16 eastern Asia; Tasmania; North America
Cupressaceae 19 130 cosmopolitan
Higher Plant Diversity
Ranunculales
69
Table 8.2 Vascular plants: a summary of systematic diversity
MAJOR GROUP (CLASS)
SUBCLASS
ORDER
FAMILY GENERA SPECIES DISTRIBUTION
Southern Hemisphere (excluding Africa) to
Araucariaceae 2 38 Indochina and the Philippines
Cephalotaxaceae 1 7 eastern Himalayas to Japan
Phyllocladaceae 1 7 Malaysia; Tasmania; New Zealand
mostly Southern Hemisphere, extending
north to Japan, Central America, and West
Podocarpaceae 6 125 Indies
Gymnosperms - Ginkgo
Ginkgoaceae 1 1 eastern China
Gymnosperms - Gnetophytes
warm temperate North and South America;
Ephedraceae 1 40 warm temperate Eurasia
tropical (Indomalaya; Fiji; northern tropical
Gnetaceae 1 30 South America; western tropical Africa)
Welwitschiaceae 1 1 southwestern Africa
Angiosperms - Dicots
Magnoliidae
Magnoliales
Winteraceae 9 100 primarily islands of southwestern Pacific
Degeneriaceae 1 1 Fiji
New Guinea; Molucca Is.; northeastern
Himantandraceae 1 1-3 Australia
Eupomatiaceae 1 2 New Guinea and eastern Australia
Austrobaileyaceae 1 1 northeastern Australia
Magnoliaceae 12 220 widespread, especially Northern Hemisphere
Lactoridaceae 1 1 San Juan Islands (Chile)
Annonaceae 130 2,300 mainly tropical
Myristicaceae 15 300 tropical
Canellaceae 6 20 tropical Africa; Madagascar; South America
Laurales
Amborellaceae 1 1 New Caledonia
New Guinea; New Caledonia; Fiji;
Trimeniaceae 2 5 southeastern Australia
tropical and subtropical, especially Southern
Monimiaceae 30-35 450 Hemisphere
Gomortegaceae 1 1 central Chile
Calycanthaceae 3 5 China; North America
Idiospermaceae 1 1 northern Australia
Lauraceae 30-50 2,000 tropical and subtropical
Hernandiaceae 4 60 tropical
Piperales
Chloranthaceae 5 TAS tropical and subtropical
eastern Asia; eastern and western North
Saururaceae = 7) America
Piperaceae 10 1,400-2,000 tropical
Aristolochiales
Aristolochiaceae 8-10 600 mainly tropical
Illiciales
Southeast Asia; southeastern United States;
IIliciaceae 1 40 Caribbean; Mexico
tropical and temperate eastern Asia;
Schisandraceae 2 50 southeastern United States
Nymphaeales
warm Asia and Australia; eastern United
Nelumbonaceae 1 2 States
Nymphaeaceae 5 50 cosmopolitan distribution
Barclayaceae 1 4 tropical Southeast Asia to New Guinea
Cabombaceae 2 8 tropical and warm temperate
Ceratophyllaceae 1 6 cosmopolitan
1. Biological Diversity
Table 8.2 Vascular plants: a summary of systematic diversity
MAJOR GROUP (CLASS)
SUBCLASS
ORDER
FAMILY GENERA SPECIES DISTRIBUTION
widespread, especially North temperate and
Ranunculaceae 50 2,000 boreal
Circaeasteraceae 2 rd Southeast Asia
widespread, especially temperate Northern
Berberidaceae 13 650 Hemisphere
Sargentodoxaceae 1 1 China, Laos, Vietnam
Lardizabalaceae 8 30 Himalayas to Southeast Asia; Chile
Menispermaceae 70 400 tropical and subtropical
Coriariaceae 1 5 disjunct in tropical America, Europe, Asia
Sabiaceae 3 60 Southeast Asia; tropical America
Papaverales
Papaveraceae 25 200 temperate & tropical Northern Hemisphere
Fumariaceae 19 400 mainly North temperate; also South Africa
Hamamelidae
Trochodendrales
Nepal; central and southeastern China;
Tetracentraceae 1 1 Burma
Trochodendraceae 1 1 Korea, Japan to Taiwan
Hamamelidales
Cercidiphyllaceae 1 2 China; Japan
Eupteleaceae 1 az Japan, China, Assam
eastern Mediterranean to Himalayas;
Platanaceae 1 6-7 Mexico to Canada
Hamamelidaceae 26 100 widespread, especially eastern Asia
Myrothamnaceae 1 2 Africa, Madagascar
Daphniphyllales
Daphniphyllaceae 1 35 Asia and Malay Archipelago
Didymelales
Didymelaceae 1 2 Madagascar
Eucommiales
Eucommiaceae 1 1 montane forests of western China
Urticales
Ulmaceae 18 150 widespread, especially Northern Hemisphere
Barbeyaceae 1 1 northeastern Africa and adjacent Arabia
Cannabaceae 2 3 North temperate
Moraceae 40 1,000 tropical and subtropical
Cecropiaceae 6 276 tropical
Urticaceae 45 700 tropical and subtropical
Leitneriales
Leitneriaceae 1 1 southeastern United States
Juglandales
widespread in Northern Hemisphere and into
Juglandaceae 7-8 60 South America
Rhoipteleaceae 1 1 southwestern China and North Vietnam
Myricales
Myricaceae 3 50 mostly temperate and subtropical
Fagales
Southwest Pacific, especially New
Balanopaceae 1 <) Caledonia
cosmopolitan, except tropical and South
Fagaceae 6-8 800 Africa
mainly temperate and cool Northern
Betulaceae 6 120 Hemisphere
Casuarinales
Casuarinaceae 1 50 Australia, Pacific islands, Asia
Caryophyllidae
Caryophyllales
Phytolaccaceae 18 125 tropical and subtropical
warm North America; Central America;
Achatocarpaceae 74 8 South America
Higher Plant Diversity
71
Table 8.2 Vascular plants: a summary of systematic diversity
MAJOR GROUP (CLASS)
SUBCLASS
ORDER
FAMILY GENERA SPECIES DISTRIBUTION
tropical and subtropical, especially New
Nyctaginaceae 30 300 World
Aizoaceae 12 2,500 South Africa; Australia
Didiereaceae 4 11 Madagascar
Cactaceae 30-200 1,000-2,000 American deserts
cosmopolitan, especially deserts and
Chenopodiaceae 100 1,500 semideserts
Amaranthaceae 65 900 tropical and subtropical
cosmopolitan, especially western North
Portulacaceae 20 500 America and Andes
Basellaceae 4 15-20 tropical and subtropical, mostly New World
Molluginaceae 13 100 tropical and subtropical, especially Africa
Caryophyllaceae 75 2,000 widespread, especially North America
Polygonales
Polygonaceae 30 1,000 mainly temperate Northern Hemisphere
Plumbaginales
Plumbaginaceae 12 400 widespread, especially Mediterranean
Dilleniidae
Dilleniales
Dilleniaceae 10 350 tropical and subtropical, especially Australia
Paeoniaceae 1 30 Eurasia, especially temperate eastern Asia
Theales
Ochnaceae 30 400 tropical, especially Brazil
Sphaerosepalaceae 2 14 Madagascar
Sarcolaenaceae 10 30 Madagascar
Dipterocarpaceae 16 600 tropical, especially rain forests of Malaysia
Caryocaraceae 2 23 tropical America, especially Amazon basin
Theaceae 40 600 tropical and subtropical
Actinidiaceae 3 300 tropical and subtropical
Scytopetalaceae 5 20 tropical western Africa
southern China to Malay peninsula and
Pentaphylacaceae 1 1 Sumatra
Malaysia; southern Venezuela (Guayana
Tetrameristaceae 2 2 Highlands)
Pellicieraceae 1 1 Costa Rica, Panama, Columbia
Oncothecaceae 1 1 New Caledonia
Marcgraviaceae 5 100 tropical America
Quiinaceae 4 40 tropical America, especially Amazon basin
Elatinaceae 2 40 tropical and subtropical
Paracryphiaceae 1 1 New Caledonia
Medusagynaceae 1 1 Seychelles
Guttiferae
(= Clusiaceae) 50 1,200 moist tropical and North temperate
Malvales
Elaeocarpaceae 10 400 tropical and subtropical
Tiliaceae 50 450 tropical and subtropical
Sterculiaceae 65 1,000 tropical and subtropical
: tropical, especially Central and South
Bombacaceae 20-30 200 America
Malvaceae 75 1,000-1,500 cosmopolitan, especially tropical
Lecythidales
tropical, especially rain forests of South
Lecythidaceae 20 400 America
Nepenthales
easter and northwestern United States;
Sarraceniaceae 3 15 northern South America
East Indies to Madagascar; to northern
Nepenthaceae 1 75 Australia and Southeast Asia
Droseraceae 4 100 temperate and tropical
Violales
1. Biological Diversity
Table 8.2 Vascular plants: a summary of systematic diversity
MAJOR GROUP (CLASS)
SUBCLASS
ORDER
FAMILY GENERA SPECIES DISTRIBUTION
Flacourtiaceae 85 800 tropical
Bixaceae 3 15 tropical
Peridiscaceae 72 2 tropical South America
Cistaceae 8 200 mostly in temperate and warm temperate
Huaceae 2 3 tropical Africa
Lacistemataceae 2 20 tropical America
Scyphostegiaceae 1 1 Borneo
Stachyuraceae 1 5-6 Himalayan region to Japan
Violaceae 16 800 cosmopolitan
Eurasia and Africa, especially Mediterranean
Tamaricaceae 45 100 region
cosmopolitan, especially Mediterranean
Frankeniaceae 3 80 region
Dioncophyllaceae 3 3 rain forests of tropical Africa
Ancistrocladaceae 1 15-20 Southeast Asia; India; tropical Africa
tropical and subtropical America and Africa;
Turneraceae 8 120 Madagascar
Malesherbiaceae 1-2 25 Andes from Chile to Peru
tropical and warm temperate, especially
Passifloraceae 16 650 tropical America and Africa
Caricaceae 4 30 tropical and subtropical America; Africa
Achariaceae 3 3 South Africa
arid parts of Mexico and southwestern
Fouquieriaceae 1 11 United States
Hoplestigmataceae 1 2 western tropical Africa
tropical and subtropical; rarely temperate or
Cucurbitaceae 90 700 cool temperate
Datiscaceae 3 a Malesia; Asia; western North America
Begoniaceae 3-5 1,020 tropical, especially northern South America
temperate and tropical North and South
Loasaceae 14 200 America
Salicales
mostly North temperate; also Australia and
Salicaceae 2 340 Malay Archipelago
Capparales
Tovariaceae 1 2 tropical America
Capparaceae 45 800 tropical and subtropical
Cruciferae cool temperate or warm temperate Northern
(= Brassicaceae) 350 3,000 and Southern Hemisphere
Moringaceae 1 10 xeric Africa; Madagascar; India
Northern Hemisphere, mostly Old World,
Resedaceae 6 70 especially Mediterranean
Batales
Gyrostemonaceae 5 17 Australia
tropical and subtropical America;
Galapagos; Hawaii; New Guinea and
Bataceae 1 7 northeastern Australia
Ericales
northern South America; Central America;
Cyrillaceae 3 14 West Indies; southeastern United States
tropical America; southeastern United
Clethraceae 1 65 States; Southeast Asia; East Indies
Grubbiaceae 1 3 South Africa (Cape Province)
cold Northern Hemisphere; southern South
Empetraceae 3 5 America; eastern United States; Europe
mostly Australia, New Zealand, and East
Epacridaceae 30 400 Indies
temperate, cool and subtropical regions;
Ericaceae 125 3,500 montane tropical
Northern Hemisphere, especially temperate
Pyrolaceae 4 45 and boreal
Higher Plant Diversity
Table 8.2 Vascular plants: a summary of systematic diversity
MAJOR GROUP (CLASS)
73
SUBCLASS
ORDER
FAMILY GENERA SPECIES DISTRIBUTION
Monotropaceae 10 12
Diapensiales
arctic & North temperate; south to
Diapensiaceae 6 18 Himalayas
Ebenales
Sapotaceae 70 800 tropical
Ebenaceae 5 450 tropical and subtropical
Styracaceae 10 150 widely disjunct in both hemispheres
Lissocarpaceae 1 2 tropical South America
tropical and subtropical America; southern
Symplocaceae 1 300-400 and eastern Asia; Australia; East Indies
Primulales
Theophrastaceae 4 100 mostly New World tropical
tropical and subtropical New and Old World;
Myrsinaceae 30 1,000 also temperate Old World
mostly temperate and cold Northern
Primulaceae 30 1,000 Hemisphere; montane tropical
Rosales
Brunelliaceae 1 50 tropical America
Connaraceae 16-24 300-400 tropical, especially Old World
Eucryphiaceae 1 6 eastern Australia; Tasmania; Chile
Southern Hemisphere, especially Australia,
New Guinea and New Caledonia; also
Cunoniaceae 25 350 Mexico and West Indies
Davidsoniaceae 1 1 northeastern Australia
Dialypetalanthaceae 1 1 Brazil
tropical and warm temperate Old World,
Pittosporaceae 9 200 especially Australia
Byblidaceae 2 4 Australia and South Africa
temperate and subtropical Northern
Hydrangeaceae 17 170 Hemisphere; southeastern Asia and Malesia
Columelliaceae 1 4 Andes, from Colombia to Bolivia
Grossulariaceae 25 350 cosmopolitan
Greyiaceae 1 3 South Africa
Bruniaceae 12 75 South Africa and Natal
tropical or subtropical forests, mostly Africa
Anisophylleaceae 4 40 and Indomalaysia; South America
Alseuosmiaceae 3 17 New Zealand and New Caledonia
cosmopolitan, except Australia and
Crassulaceae 25 900 Polynesia
Cephalotaceae 1 1 southwestern Australia
cosmopolitan, especially temperate and cold
Saxifragaceae 40 700 Northern Hemisphere
cosmopolitan, especially temperate and
Rosaceae 100 3,000 subtropical Northern Hemisphere
deserts in Africa, across Middle East to
Neuradaceae 3 10 India
arid western United States and adjacent
Crossosomataceae 3 10 Mexico
Chrysobalanaceae 17 450 pantropical, especially New World
Surianaceae 4 6 Australia and tropical maritime
Rhabdodendraceae 1 3 tropical South America
Fabales
Leguminosae cosmopolitan, especially tropical and
(= Fabaceae) 590 14,200 subtropical
Proteales
temperate and subtropical Northern
Hemisphere, to tropical Asia and northern
Elaeagnaceae 3 50 Australia
1. Biological Diversity
Table 8.2 Vascular plants: a summary of systematic diversity
MAJOR GROUP (CLASS)
SUBCLASS
ORDER
FAMILY
Proteaceae
Podostemales
Podostemaceae
Haloragales
Haloragaceae
Gunneraceae
Myrtales
Sonneratiaceae
Lythraceae
Penaeaceae
Crypteroniaceae
Thymelaeaceae
Trapaceae
Myrtaceae
Punicaceae
Onagraceae
Oliniaceae
Melastomataceae
Combretaceae
Rhizophorales
Rhizophoraceae
Cornales
Alangiaceae
Nyssaceae
Cornaceae
Garryaceae
Santalales
Medusandraceae
Olacaceae
Dipentodontaceae
Opiliaceae
Santalaceae
Misodendraceae
Loranthaceae
Viscaceae
Eremolepidaceae
Balanophoraceae
Rafflesiales
Hydnoraceae
Mitrastemonaceae
Rafflesiaceae
Celastrales
Geissolomataceae
Celastraceae
Hippocrateaceae
Salvadoraceae
Stackhousiaceae
Aquifoliaceae
Icacinaceae
GENERA
75
40
2-13
50
SPECIES
1,000
200
20
7-8
20-25
320-420
400
74
DISTRIBUTION
tropical and subtropical, especially warmer
Southern Hemisphere
mostly tropical, especially Asia and America
cosmopolitan, especially Southern
Hemisphere
Southern Hemisphere to southern Mexico
Old World tropical
mainly tropical; also temperate
Cape Province (South Africa)
India, Philippines, Malay Archipelago
cosmopolitan
tropical and subtropical Africa and Eurasia
tropical and subtropical; temperate Australia
Balkans to northern India; Socotra
temperate and subtropical, especially New
World
tropical and southern Africa; St Helena
tropical and subtropical, especially South
America
tropical and subtropical, especially Africa
tropical and subtropical
eastern and tropical Asia; eastern Australia;
Pacific islands; Madagascar; western Africa
eastern North America; eastern Asia; Pacific
islands; China
North temperate; irregularly tropical and
South temperate
western North and Central America, from
Washington to Panama
rainforests of tropical western Africa
tropical and subtropical
southern China and Burma
tropical and subtropical
nearly cosmopolitan, especially arid climates
temperate South America
mostly tropical and subtropical
cosmopolitan, especially tropical
tropical America
tropical and subtropical
drier parts of Africa, Madagascar
Borneo and Sumatra to Indochina and
Japan; Mexico and Central America
tropical and subtropical
South Africa (Cape Province)
pantropical, some in temperate regions
tropical
Africa; Madagascar; India; Sri Lanka;
Southeast Asia
Australia and New Zealand; southwestern
Pacific
more or less cosmopolitan
pantropical
Table 8.2
MAJOR GROUP (CLASS)
SUBCLASS
ORDER
FAMILY GENERA
Aextoxicaceae 1
Cardiopteridaceae 1
Corynocarpaceae 1
Dichapetalaceae 3
Euphorbiales
Buxaceae 5
Simmondsiaceae 1
Pandaceae 3
Euphorbiaceae 300
Rhamnales
Rhamnaceae 55
Leeaceae 1
Vitaceae 11
Linales
Erythroxylaceae 4
Humiriaceae 8
Ixonanthaceae 5
Hugoniaceae 7
Linaceae 6
Polygalales
Malpighiaceae 60
Vochysiaceae 7
Trigoniaceae 3
Tremandraceae 3
Polygalaceae 12
Xanthophyllaceae 1
Krameriaceae
Sapindales
Staphyleaceae 5
Melianthaceae 2
Bretschneideraceae 1
Akaniaceae 1
Sapindaceae 140
Hippocastanaceae 2
Aceraceae 2
Burseraceae 16-20
Anacardiaceae 60-80
Julianiaceae 2
Simaroubaceae 25
Cneoraceae 1
Meliaceae 51
Rutaceae 150
Zygophyllaceae 30
Geraniales
SPECIES
1,200
200
750
75
Higher Plant Diversity
Vascular plants: a summary of systematic diversity
DISTRIBUTION
Chile
Asia to New Guinea and Australia
New Zealand; northeastern Australia; New
Guinea
pantropical, mainly Africa
nearly cosmopolitan
western United States and Mexico
Africa, Asia, New Guinea
cosmopolitan, especially tropical and
subtropical
cosmopolitan, especially tropical and
subtropical
pantropical
tropical and subtropical; a few in temperate
regions
pantropical, especially New World
mainly tropical South America, with one
species in Africa
pantropical
tropical
widespread, especially temperate and
subtropical
tropical and subtropical, especially South
America
mostly tropical America, 1 in Africa
subtropical in moist lowland forests
Australia and Tasmania
nearly cosmopolitan
Indomalaysian region
Argentina and Chile, mainly in dry regions
Americas, Eurasia, Malay Archipelago
Africa
mountains of western and southwestern
China
eastern Australia
tropical and subtropical; some in temperate
regions
North America to northern South America;
Europe; Southeast Asia
temperate and subtropical, especially
Malesia; China
pantropical, especially tropical America and
Northeast Africa
mainly pantropical, some in temperate
regions
tropical America (Central America, Peru)
pantropical, some in warm temperate
regions
Mediterranean, Canary Is., Cuba
tropical and subtropical; some in temperate
regions
nearly cosmopolitan, especially South Africa
and Australia
mostly arid tropical and subtropical,
sometimes in saline habitats
1. Biological Diversity
Table 8.2
MAJOR GROUP (CLASS)
SUBCLASS
ORDER
FAMILY
Oxalidaceae
Geraniaceae
Limnanthaceae
Tropaeolaceae
Balsaminaceae
Apiales
Araliaceae
Umbelliferae
(= Apiaceae)
Asteridae
Gentianales
Loganiaceae
Retziaceae
Gentianaceae
Saccifoliaceae
Apocynaceae
Asclepiadaceae
Solanales
Nolanaceae
Duckeodendraceae
Solanaceae
Convolvulaceae
Cuscutaceae
Menyanthaceae
Polemoniaceae
Hydrophyllaceae
Lamiales
Lennoaceae
Boraginaceae
Verbenaceae
Labiatae
(= Lamiaceae)
Callitrichales
Hippuridaceae
Callitrichaceae
Hydrostachyaceae
Plantaginales
Plantaginaceae
Scrophulariales
Buddlejaceae
300
250
85
50
18
20
10
SPECIES
900
700
11
92
450
700
3,000
250
45
76
Vascular plants: a summary of systematic diversity
DISTRIBUTION
tropical and subtropical; some in temperate
regions
temperate and warm temperate regions;
some tropical
temperate North America
Mexico to Chile (in mountains), Patagonia
tropical Asia and Africa, some in temperate
regions; India to Java
tropical and subtropical; some in temperate
regions
nearly cosmopolitan, especially North
temperate regions and tropical mountains
tropical and subtropical; relatively few
species in temperate regions
Cape Province of South Africa
cosmopolitan, especially temperate and
subtropical regions and tropical mountains
southern Venezuela
tropical and subtropical; relatively few
species in temperate regions
tropicals and subtropical, especially Africa,
with relatively few species in temperate
regions
northern Chile and southern Peru, often
along the seashore
Amazon basin of Brazil
nearly cosmopolitan, especially tropical
South America
nearly cosmopolitan, especially tropical and
subtropical
nearly cosmopolitan, especially warmer
parts of New World
cosmopolitan
North temperate (Eurasia, Alaska to western
South America), especially temperate North
America
wide-ranging, especially dry western United
States
New World from southwestern United
States to Colombia and Venezuela
cosmopolitan, especially western North
America and Mediterranean region; east into
Asia
pantropical, with only a few species in
temperate regions
cosmopolitan, especially Mediterranean
region and into central Asia
temperate and boreal Northern Hemisphere;
Australia; southern South America
nearly cosmopolitan
Madagascar; tropical and southern Africa
cosmopolitan
mainly tropical and subtropical
Table 8.2
MAJOR GROUP (CLASS)
SUBCLASS
ORDER
FAMILY
Oleaceae
Scrophulariaceae
Globulariaceae
Myoporaceae
Orobanchaceae
Gesneriaceae
Acanthaceae
Pedaliaceae
Bignoniaceae
Mendonciaceae
Lentibulariaceae
Campanulales
Pentaphragmataceae
Sphenocleaceae
Campanulaceae
Stylidiaceae
Donatiaceae
Brunoniaceae
Goodeniaceae
Rubiales
Rubiaceae
Theligonaceae
Dipsacales
Caprifoliaceae
Adoxaceae
Valerianaceae
Dipsacaceae
Calycerales
Calyceraceae
Asterales
Compositae
(= Asteraceae)
Angiosperms - Monocots
Alismatidae
Alismatales
Butomaceae
Limnocharitaceae
Alismataceae
Hydrocharitales
Hydrocharitaceae
Najadales
Aponogetonaceae
Scheuchzeriaceae
GENERA
30
190
10
3-4
120
450
1,100
12
15
SPECIES
600
4,000
300
125
17
300
270
20,000
77
Higher Plant Diversity
Vascular plants: a summary of systematic diversity
DISTRIBUTION
nearly cosmopolitan, especially Asia and
Malesia
cosmopolitan, especially temperate regions
and tropical mountains
Africa; Madagascar; Europe; western Asia
Australia; Asia; Pacific islands; West Indies;
northern South America
150
pantropical, with a few species in temperate
regions
tropical, with only a few species in
temperate regions
mostly tropical, especially along seacoast or
in arid regions, with only a few species in
temperate climates
mainly tropical, especially tropical America
South America; tropical Africa; Madagascar
cosmopolitan
Southeast Asia and nearby Pacific islands
pantropical; western Africa
cosmopolitan
Australasia; south and Southeast Asia;
southernmost South America
southern South America; New Zealand;
Tasmania
Australia
primarily Australia; also New Zealand,
Japan, and tropical and subtropical Old and
New World
cosmopolitan, especially tropical and
subtropical
temperate eastern Asia to Mediterranean
region and Canary Islands
mostly North temperate and boreal regions;
also tropical mountains
circumboreal
nearly cosmopolitan, especially North
temperate regions and Andes
Eurasia and Africa, especially Mediterranean
region
Central and South America
cosmopolitan, especially temperate and
subtropical regions
temperate Eurasia
tropical and subtropical
cosmopolitan, especially Northern
Hemisphere
cosmopolitan
Old World tropical to South Africa
cool Northern Hemisphere
1. Biological Diversity
Table 8.2
MAJOR GROUP (CLASS)
SUBCLASS
ORDER
FAMILY
Juncaginaceae
Potamogetonaceae
Ruppiaceae
Najadaceae
Zannichelliaceae
Posidoniaceae
Cymodoceaceae
Zosteraceae
Triuridales
Triuridaceae
Petrosaviaceae
Arecidae
Arecales
Palmae
(= Arecaceae)
Cyclanthales
Cyclanthaceae
Pandanales
Pandanaceae
Arales
Araceae
Lemnaceae
Commelinidae
Commelinales
Rapateaceae
Xyridaceae
Mayacaceae
Commelinaceae
Eriocaulales
Eriocaulaceae
Restionales
Flagellariaceae
Restionaceae
Joinvilleaceae
Centrolepidaceae
Juncales
Juncaceae
Thurniaceae
Cyperales
Cyperaceae
Gramineae
(= Poaceae)
Hydatellales
Hydatellaceae
Typhales
Sparganiaceae
Typhaceae
GENERA
ou-poe- seu
16
70
500
SPECIES
682-782
1,800
31
35
10
78
Vascular plants: a summary of systematic diversity
DISTRIBUTION
temperate and cold Northern and Southern
Hemisphere
cosmopolitan
temperate and subtropical
cosmopolitan
cosmopolitan
Mediterranean, Australia
tropical and subtropical seacoasts
subarctic, temperate, subtropical seacoasts
tropical and subtropical
southern China and southern Japan to
Malay Peninsula and Borneo
tropical and warm temperate
tropical America
Old World, especially tropical (Malesia)
mostly tropical and subtropical
cosmopolitan
tropical South America, with one species in
tropical western Africa
tropical and subtropical; a few species in
temperate region
tropical western Africa; tropical and warm
temperate America
tropical and subtropical
tropical and subtropical, with a few species
in temperate regions
Old World tropical
widely distributed in Southern Hemisphere,
especially Australia and South Africa
Pacific Islands
Australia; Southeast Asia; Pacific Islands;
southernmost South America; mostly in
nutrient-poor soils
temperate or cold regions, or montane
tropical
Amazon basin and Guayana
cosmopolitan, most abundant in temperate
regions
cosmopolitan, especially tropical and North
temperate semi-arid regions with seasonal
rainfall
Australia, New Zealand, Tasmania
chiefly North temperate regions, to Australia
and New Zealand
cosmopolitan
Table 8.2
MAJOR GROUP (CLASS)
SUBCLASS
ORDER
FAMILY
Zingiberidae
Bromeliales
Bromeliaceae
Zingiberales
Liliidae
Strelitziaceae
Heliconiaceae
Musaceae
Lowiaceae
Zingiberaceae
Costaceae
Cannaceae
Marantaceae
Liliales
Philydraceae
Pontederiaceae
Haemodoraceae
Cyanastraceae
Liliaceae
lridaceae
Velloziaceae
Aloeaceae
Agavaceae
Xanthorrhoeaceae
Hanguanaceae
Taccaceae
Stemonaceae
Smilacaceae
Dioscoreaceae
Orchidales
Burmanniaceae
Geosiridaceae
Corsiaceae
Orchidaceae
GENERA
47
30
280
18
12
20
1
2
SPECIES
2,000
100
800-1,000 25,000-35,000
Higher Plant Diversity
Vascular plants: a summary of systematic diversity
DISTRIBUTION
New World, except one species in western
tropical Africa
tropical
tropical and subtropical South and Central
America; one species widespread in
southwestern Pacific islands
tropical and subtropical Old World
southern China; Malay Peninsula; Pacific
islands
tropical regions, especially southern and
Southeast Asia
pantropical, especially New World
tropical and subtropical New World
pantropical, especially New World
Australia; western Pacific islands to Japan
and mainland Southeast Asia
tropical and subtropical; into North
temperate regions
mostly Southern Hemisphere, but reaching
northern United States
forests of tropical Africa
widespread, especially dry, temperate to
subtropical regions
cosmopolitan, especially South Africa
South America; Africa; Madagascar;
southern Arabia
Africa, Madagascar, Arabia, nearby islands;
especially South Africa
warm, mostly arid regions of New and Old
Worlds; a few in distinctly temperate
climates
Australia; Tasmania; New Guinea; New
Caledonia
Malesia; Sri Lanka
pantropical, especially Southwest Asia and
Polynesia
eastern Asia; Malesia; northern Australia;
southeastern United States
tropical and subtropical, especially Southern
Hemisphere; also in parts of North
temperate region
tropical and subtropical, with a few species
in North temperate region
pantropical, with a few species in temperate
regions
Madagascar and other Indian Ocean islands
New Guinea, Chile
cosmopolitan
Sources: Flowering plant information modified from Cronquist, A. 1981. An Integrated System of Classification of Flowering Plants. Columbia
Univ. Press. 1262 pp.; other information from Airy Shaw, H.K. (ed.). A Dictionary of the Flowering Plants and Ferns. Eighth Edition. Cambridge
Univ. Press. 1245 pp.; and other sources.
719
1. Biological Diversity
Table 8.3 Species richness and endemism: higher plants
FLOWERING GYMNO-— FERNS NUMBER OF % ESTIMATE/ COMPLETION DATE
PLANTS SPERMS ENDEMICS ENDEMISM COUNT
ASIA
Afghanstan 3,500 = = = [30-35%] e2 2 1989-91
Bahrain 195 1 1 0 0.0 c 1 1991
Bangladesh 5,000 = - - = e2 2 1972
Bhutan 5,446 22 - 50-100 14 e1 3 1991
British Indian Ocean Territory 100 1 = i?) 0.0 e1 1 1971
Brunei 3,000 28 = 7 0.2 e2 5 1990
Cambodia = = = = = = = =
China 30,000 200 2,000 18,000 55.9 e2 3 1991
Cyprus 1,650 12 20 88 5.2 c 1 1977-85
Hong Kong 1,800 4 180 25 1.3 e2 2 1978-91
India 15,000 - 1,000 5,000 31.3 e2 2 1983-84
Indonesia 20,000 - 2,500 15,000 66.7 e3 4 1991
Iran, Islamic Rep 6,500 33 - - [30-35%] e2 1 1989-91
lraq 2,914 7 16 190 6.5 c 1 1966-86
Israel 2,294 8 15 155 6.7 c 1 1982-84
Japan 4,700 42 630 2,000 37.2 c 1 1987
Jordan 2,200 6 6 = = c 2 1982-85
Korea, Dem People’s Rep {2,898 = = 107 {14.0 {c = {1976-83
Korea, Rep {2,898 - - 224 {14.0 {ce - {1976-83
Kuwait 234 1 1 0 0.0 c 1 1991
Laos = = = = = = = =
Lebanon 2,000 12 40 = [10%] e3 2 1984-91
Malaysia 12,000 = 500 = = e3 3 1991
Maldives 260 2 15 5 1.8 c 1 1983
Mongolia 2,272 = =. 229 10.1 c 1 1984
Myanmar 7,000 - = 1,071 15.3 e2 4 1961
Nepal 6,500 23 450 315 4.5 c 2 1978-82
Oman 1,018 3 14 74 7A c 1 1991
Pakistan 4,917 21 - 372 75 e1 2 1986
Philippines 8,000 31 900 3,500 39.3 e2 3 1982-91
Qatar 220 1 te) ts) 0.0 c 1 1991
Saudi Arabia 1,729 8 22 34 1.9 c 2 1991
Singapore 2,000 2 166 1 0.1 e1 1 1989-91
Sri Lanka 2,900 = 314 900 28.0 c 2 1982-83
Syria 2,000 12 40 = [10%] e3 2 1984-91
Taiwan 2,983 20 565 = [25%] c 1 1982-91
Thailand 12,000 25 600 = = e2 3 1979-85
Turkey 8,472 22 85 2,651 30.9 c 1 1988
United Arab Emirates 340 2 5 ts) = c 1 1991
Viet Nam = = = = = = = =
Yemen, People’s Dem Rep! 1,373 3 41 58 41 c 2 1991
Yemen, Arab Rep! 959 1 14 77 7.9 c 1 1991
USSR?
22,000 74 207 - - e1 2 1991
EUROPE
Albania 2,965 21 45 24 0.8 c 2 1980-88
Andorra 980 6 26 oO 0.0 e1 1 1981
Austria 2,850—3,050 12 66 35 1.2 el 1 1978-91
Belgium 1,250—1,550 2 50 1 0.1 e1 1 1978-83
Bulgaria 3,505 15 52 320 9.0 c 1 1991
Czechoslovakia 2,507 11 72 62 2.4 c 1 1991
Denmark 1,000—1,400 2 50 1 0.1 e1 1 1984-91
Faeroe Islands 236 1 25 1 0.4 c 1 1991
Finland 1,040 4 58 0 0.0 c 1 1988
France 4,500 20 110 133 2.9 c 1 1991
Germany 2,600 10 72 6 0.2 e1 1 1984-91
Greece 4,900 21 71 742 14.9 e1 2 1989
Hungary 2,148 8 58 38 17 c 1 1991
Iceland 340 1 36 1 0.3 e1 1 1984-91
lreland 892 2 56 i*} 0.0 c 1 1991
Italy 5,463 29 106 712 12.7 c 1 1982
Liechtenstein 1,400 10 = ts) 0.0 e2 2 1977
Luxembourg 1,200 4 42 0 0.0 e1 1 1984-91
Malta 900 3 11 5 0.5 e1 1 1984
Monaco = 4 18 () 0.0 = = 1973
Netherlands 1,170 3 48 0 0.0 c 1 1991
Norwai 1,550—1,750 4 61 1 01 e1 1 1978-91
Polan 2,200—2,400 10 62 3 0.1 e1 1 1978-91
Portugal® 2,400—2,600 8 65 150 5.8 e1 1 1978-91
Romania 3,000—3,350 11 62 41 1.3 e2 2 1977-78
San Marino = = = to) 0.0 = = 1991
Spain* 4,916 18 114 941 18.6 c 2 1984-91
Sweden 1,550-1,750 4 60 1 0.1 e1 1 1978-91
Switzerland 2,927 16 87 1 0.1 c 1 1989
United Kingdom 1,550 3 70 16 1.0 e1 1 1991
Vatican City - = - to) 0.0 = = 1991
Yugoslavia 5,250 23 78 137 2.6 e2 2 1978-91
80
Higher Plant Diversity
Table 8.3 Species richness and endemism: higher plants (continued)
FLOWERING GYMNO—- FERNS NUMBER OF % ESTIMATE/ COMPLETION DATE
PLANTS SPERMS ENDEMICS ENDEMISM COUNT
NORTH AND CENTRAL AMERICA
Anguilla 321 = 0 1 0.3 c = 1991
Antigua and Barbuda 766 1 33 = [0.7%] c 1 1938-91
Aruba 460 = = 25 5.4 c = 1991
Bahamas 1,172 3 * 43 115 9.4 c 1 1982-91
Barbados 542 = 30 5 0.8 c 2 1984-91
Belize 2,500—3,000 10 134 150 5.2 e2 2 1989-91
Bermuda 147 0 20 15 9.0 c 1 1991
Canada 2,920 33 65 147 4.9 c 1 1967-91
Cayman Islands 518 1 20 19 3.4 c 1 1984
Costa Rica 10,000—12,000 9 1,000 1,800 15.0 e2 3 1989-91
Cuba 5,996 23 495 3,229 49.6 c 2 1991
Dominica 1,127 1 197 11 0.8 c 1 1991
Dominican Republic {5,000 {7 {650 {1,800 = c {2 1984-91
El Salvador 2,500 8 400 17 0.6 e2 3 1989-91
Greenland (Denmark) 497 1 31 0 0.0 c 1 1978
Grenada 919 1 148 4 0.4 e3 2 1979-91
Guadeloupe {1670 1 261 26 1.6 c 1 1991
Guatemala 8,000 29 {652 1,171 13.5 e2 3 1989-91
Haiti {5,000 {7 {650 {1,800 31.8 c {2 1984-91
Honduras 5,000 30 325 148 28 e2 3 1978-91
Jamaica 2,746 4 558 906 27.4 c 2 1991
Martinique {1670 1 259 30 1.9 c 1 1979-91
Mexico 20,000—30,000 71 1,000 3,624 13.9 e2 3 1984-91
Montserrat 554 = 117 2 0.3 c 1 1991
Netherlands Antilles = = = = = = - =
Nicaragua 7,000 - 500 57 0.8 e2 3 1989-91
Panama 9,000 12 577 1,222 12.7 e2 3 1989-91
Puerto Rico 2,128 1 364 235 9.4 e2 2 1982-91
St Kitts and Nevis 533 = 122 = = c 1 1979
St Lucia 909 = 118 11 1.1 c = 1991
St Vincent and the Grenadines 1,000 1 165 = = e3 3 1979-91
Trinidad and Tobago 2,132 = 289 226 9.3 e2 1 1981-91
Turks and Caicos Islands 440 1 7 9 2.0 c 1 1982
United States 18,956 113 404 4,036 20.7 c 2 1978-91
Virgin Islands (British) = = = = = = = -
Virgin Islands (US) = = = = = - = —
SOUTH AMERICA
Argentina 9,000 13 359 - [25-30%] e1 2 1984-91
Bolivia 15,000—18,000 = = = = e3 5 1989
Brazil 55,000 - - = - e3 4 1979
Chile 4,750—5,500 17 150 2,698 51.1 e1 2 1983-91
Colombia 35,000 = = 1,500 43 e2 4 1989
Ecuador 16,500—20,000 - 1,100 4,000 20.7 e2 4 1986-91
French Guiana 5,000 = 318 = - e1 2 1991
Guyana 6,000 = = = = e3 2 1991
Paraguay 7,000—8,000 = = = = e3 4 1985
Peru 13,000 11 1,000 = = e2 4 1984-91
Suriname 4,500 2 293 = = e1 2 1978-91
Uruguay = 2 81 = = c - 1991
Venezuela 15,000—25,000 14 1,059 8,000 38.0 e3 4 1979-91
OCEANIA
American Samoa 328 0 125 10 2.2 c 1 1991
Australia 15,000 - = = [80%] c 3 1990
Cook Islands 184 te) 100 3 141 c 1 1991
Fiji 1,307 11 310 812 49.9 c 1 1991
French Polynesia = = = = = = = =
Guam 330 = = = [69%] = 1 1970
Kiribati 60 0 = 2 3.3 e2 1 1973-74
Marshall Islands 100 1 10 4 3.6 e2 2 1960-82
Micronesia, Federated States of - = = = = = = =
Nauru 50 L*} 4 1 1.9 e2 2 1982
New Caledonia 2,750 44 300 2,480 80.2 c 2 1991
New Zealand 2,160 22 189 1,942 81.9 c 1 1991
Niue 150 te) 28 1 0.6 rc 1 1991
North Marianas Islands 250 1 64 81 25.7 e2 3 1978-82
Palau = = = = = = = =
Papua New Guinea 10,000 44 1,500 = [55%] e2 4 1979-91
Pitcairn Islands 56 0 20 14 18.4 c 2 1960-83
Solomon Islands 2,780 22 370 30 0.9 e1 3 1991
Tokelau 26 (0) 6 0 0.0 c 1 1991
Tonga 360 1 102 25 5.4 c 1 1991
Tuvalu S - - = = = = =
Vanuatu 1,000 - - 50 5.0 e1 1 1978
Wallis and Futuna Islands 250 a = 5 2.0 e1 1 1983
Western Samoa 493 0 200 117 16.9 c 1 1991
81
1. Biological Diversity
Table 8.3 Species richness and endemism: higher plants (continued)
FLOWERING GYMNO-— FERNS NUMBER OF % ESTIMATE/ COMPLETION DATE
PLANTS SPERMS ENDEMICS ENDEMISM COUNT
ANTARCTICA
Antarctica 41 0 11 11 21.2 c 1 1990
Falkland Islands (Malvinas) 146 0 19 14 8.5 c 1 1991
French Southern Territories 30 te) 20 11 22.0 c 1 1990
AFRICA
Algeria 3,100 18 46 250 7.9 e1 2 1975-84
Angola 5,000 = 185 1,260 24.3 e2 3 1991
Benin {3050 {1 {200 t') 0.0 c 2 1901
Botswana - 0 15 17 = c 3 1970-78
Burkina Faso 1,100 0 = (+) 0.0 e3 3 1954-85
Burundi 2,500 - - - = e2 2 1901
Cameroon 8,000 3 257 156 1.9 e2 3 1964-83
Cape Verde 740 0 34 86 114 c 1 1985
Central African Rep 3,600 2 - 100 2.8 e3 3 1958
Chad 1,600 - = = = e1 1 1991
Comoros 660 1 60 136 18.9 c 2 1917
Congo 4,350 7 = - [5—10%] e2 2 1988-91
Cote d'Ivoire 3,517 (0) 143 62 17 c 2 1985
Djibouti 635 2 4 2 0.3 c 1 1989
Egypt 2,066 4 6 70 3.4 c 1 1974-84
Equatorial Guinea 3,000 0 250 66 2.0 e3 4 1991
Ethiopia 6,000-—7,000 3 100 600-1400 15.1 e2 4 1989
Gabon 6,000—7,000 1 150 = [5-—10%] e2 4 1991
Gambia 966 0 8 () 0.0 c 1 1991
Ghana 3,600 1 124 43 1.2 e2 2 1991
Guinea 3,000 t) - 88 2.9 e3 3 1991
Guinea-Bissau 1,000 t) = 12 1.2 e2 2 1991
Kenya 6,000 6 500 265 41 e2 2 1984
Lesotho 1,576 0 15 2 0.1 c 1 1971-75
Liberia 2,200 0 = 103 47 e3 4 1991
Libya 1,800 10 15 134 7.3 c 1 1975-84
Madagascar 8,000—10,000 5 500 5,000—8,000 68.4 e3 4 1987
Malawi 3,600 4 161 49 1.3 e2 2 1970-75
Mali 1,741 0 = 11 0.6 c 1 1991
Mauritania 1,100 (°) = = = e2 1 1976
Mauritius 700 (°) 178 329 37.5 e1 2 1978-91
Mayotte = - = = = = = -
Morocco 3,600 19 56 600-650 17.0 el 2 1975-84
Mozambique, People’s Rep 5,500 9 183 219 3.8 e1 2 1960-70
Namibia 3,128 1 45 = = c 1 1976
Niger 1,170 i?) 8 0 0.0 c 1 1983
Nigeria 4,614 1 100 205 43 e2 2 1991
Reunion 750 i) 240 175 17.7 e1 1 1991
Rwanda 2,288 2 = 26 11 c 2 1978-88
Saint Helena 50 (°) 24 59 79.7 c 1 1991
Sao Tome and Principe 744 1 150 134 15.0 c 1 1973
Senegal 2,062 (0) 24 26 1.2 c 1 1973
Seychelles 1,139 1 500 250 15.2 c 2 1989-91
Sierra Leone 1,700—2,480 0 = 74 3.5 e2 1 1962-91
Somalia 3,000 2 26 500 16.5 e2 3 1991
South Africa 2,300 40 380 = [70-80%] e2 1 1984
Sudan 3,132 5 - 50 1.6 c 2 1952-56
Swaziland 2,636 8 71 4 0.1 c 1 1983
Tanzania 10,000 8 = 1,122 11.2 e2 1 1968
Togo _ {3050 {1 _{200 (e) 0.0 c 2 1991
Tunisia 2,150 10 36 - - e1 1 1976-84
Uganda 5,000 6 400 30 0.6 e2 2 1984
Western Sahara 330 = = = = e2 2 1976
Zaire 11,000 iz, = 3,200 29.1 e2 5 1991
Zambia 4,600 1 146 211 44 e2 3 1960-70
Zimbabwe 4,200 6 234 95 21 e2 2 1970-75
Notes: { Indicates figure is a combined total with another country. This applies to both Korean nations (flowering plants); Guadeloupe and Martinique
Heserine plants); Guatemala and Belize (ferns) Benin and Togo (all plants); Dominican Rep. and Haiti (data for Hispaniola only). % endemism: calculated
rom data unless in square brackets. Estimate/count: c count; el approxiante count; e2 extrapolation; e3 estimate on basis of any available information and
comparable floras. Completion: Percentage of flora still to be described. 1: <5%(+/—known). 2:5-10%. 3: 10-15%. 4: 15-20%. 5; >20%. Date: date of
information. — no data available. ' no data available for the new combined Yemen Republic. ? USSR: covers the former Union of Soviet Socialist Republics
* Portugal: data include the Azores. * Spain: data do not include the Canary Islands
Table compiled for WCMC by John Akeroyd.
82
Higher Plant Diversity
Figure 8.1 Percent endemism of country floras
66-S [fF
Gy Ol | 4
(o1az ‘aiodeBuls ‘puejiazyims +)
eyep ou (frame
wisiwiapua %
1. Biological Diversity
2 The 25 most plant-rich countries
0 Ht PO
v5
ace
Country
oc cn? yo ye 0 VE ad? cd? eo go aor 4a? we
sajoeds jo sOqUINN
84
Higher Plant Diversity
25,000
Number of species
a 8
8 3
8
5,000
PAE SI LEE ‘ a PEP IP OLLS
C ae
ountry
5,000
:
Number of species
c*)
3
8
SIM Yi ltddlf dipped
Country
85
1. Biological Diversity
Figure 8.5 Flowering plant richness: North and Central America
25,000
S a 8
8 8 8
Number of species
8
CUE UGG GY
‘ &
Country
>
Figure 8.6 Flowering plant richness: South America and Antarctica
40,000
8
8
Number of species
8
8
| MIOGELLGL cam
&
86
Figure 8.7
Number of species
Higher Plant Diversity
Flowering plant richness: Oceania including Australia
S Yip Ul gpl PY lies
Number of species
8 Flowering plant richness: Africa and Madagascar
= ue Qe >
Country oe :
YTS. GM Y LU EY!
Country
87
1. Biological Diversity
9. NEMATODES
The phylum Nematoda includes a very large number of
very small worm-like animals which have a great impact on
humans, often directly deleterious, as with many parasitic
forms, but also with an important role in decomposition and
nutrient cycling. The group contains a large number of
described species, but the true proportion of the world’s
species that are nematodes is suspected of being very large
indeed. This section is intended to introduce some features
of nematodes important in the context of biological
diversity.
NEMATODE DIVERSITY
More than 15,000 species have been described and the total
number of species has been estimated at between 500,000
species (Poinar, 1983) and around one million (J.
Lambshead, pers comm.). Nematodes show a wide range of
life histories, from the entirely free-living to almost totally
parasitic in plants and many kinds of animals. The parasitic
forms which afflict humans, domesticated animals and
plants are among the best-studied species. Anderson (1984)
showed that approximately a third of known nematode
genera are parasitic on vertebrates (Table 9.1).
Of the non-parasitic forms, those feeding on micro-
organisms (especially bacteria) can be described as
microbotrophic, and those that feed on multicellular
metazoan organisms are described as predaceous. All others
are described as parasitic on plants and fungi, invertebrates
or vertebrates (Poinar, 1983).
Nematodes are usually long and cylindrical in shape (giving
rise to the common name ‘roundworms’) and their cuticle
is of a type of secreted collagen thought to be peculiar to
nematodes. Uniquely, muscle-nerve links arise during
development from the muscle not the nerve, as is usually
the case (Barnes, 1980). Nematodes have a relatively
complicated reproductive system and lack dispersive larvae.
These features might be implicated in the high species
richness of the group (J. Lambshead, pers. comm.). Body
length varies enormously. One of the smallest known
marine nematodes, Greeffiella minutum, is only 82um long;
however, the largest nematode known, Placentonema
gigantissima, which is parasitic in the placenta of the sperm
whale has been recorded at over 8m (Poinar, 1983).
Taxonomic procedures are difficult because of the small
size of many nematode species. There have been several
major taxonomic reviews over the last few decades.
Table 9.1
different habitats
HABITAT
Marine and freshwater
Soil
Plant (parasitic)
Invertebrate (parasitic)
Vertebrate (parasitic)
TOTALS
Classification is almost entirely based on morphological
characteristics visible under a compound microscope
(Poinar, 1983). Many species, especially those with
parasitic relationships with other organisms, cannot be kept
in culture and thus are not amenable to biochemical or
genetic study. Scientists of different disciplines frequently
work independently of each other, resulting in confusing
taxonomic revisions.
Estimates of the total number of nematode species vary
greatly, current figures ranging from 500,000 to around one
million. Recent work on species diversity in the meiofauna
of deep-sea benthic samples has found very high diversity
in each sample. However, taxonomic problems and the
sheer number of organisms involved means that the species
similarity between samples is still unresolved (J.
Lambshead, pers. comm.). If many of these samples
constitute separate species, nematodes may approach or
even exceed the insects in species richness.
Microbotrophic nematodes
The microbotrophic nematodes, especially some marine
forms, are generally thought to represent the most primitive
organisms in the phylum, although there is an alternative
hypothesis that extant microbotrophsare secondarily derived
from parasitic forms (Poinar, 1983). It is difficult to
elucidate the evolutionary history of a group which leaves
few fossil remains but it is thought that microbotrophic
nematodes were probably well represented in the Cambrian
period, c. 600 million years ago.
Microbotrophic nematodes are one of the most widespread
and abundant animal groups known. Wherever a suitable
food source exists they are found, even under extreme
conditions such as hot sulphur springs or polar ice. Because
of their relatively small size (although some grow to over
10mm, most cannot be seen with the naked eye) they tend
to go unnoticed even though present in great numbers. For
example, about 90,000 nematodes of several different
species have been found in a single rotting apple in an
orchard and about 50,000 nematodes of at least eight
different species have been reported from a single fig
(Barnes, 1980).
These nematodes can be divided into three groups - marine,
freshwater and terrestrial - although even the so called
terrestrial species are dependent upon the water film around
soil particles and in interstitial spaces. Those species which
Approximate numbers of nematode families and genera known from
FAMILIES GENERA
41 730
64 429
26 166
42 187
83 759
256 2271
Source: Anderson, R.V. 1984. The origins of zooparasitic nematodes. Canadian Journal of Zoology, 62:317-28.
live in environments with only a periodic water supply,
such as deserts, survive mostly as inactive larvae and only
emerge when water is present.
Marine species live in bottom sediments of many habitats
from sandy shores and salt-marshes to ocean trenches and
have been reported in numbers ranging from 100,000 to 10
million individuals per m? (Poinar, 1983). Thus they are the
most important metazoan element of the meiofauna in all
samples. Samples reported by Nicholas (1984) taken at
various depths down to about 400m show a range in
number of species from 3 to 125 per site and a range in
densities of 110,000 to 5,261,000 animals per m*. These
samples were derived from sediments, algae, shells and
rocks, where bacteria and other micro-organisms flourish.
In one study of deep-sea nematodes, examination of 216
individuals yielded a total of 148 species (J. Lambshead,
pers. comm.).
Several groups of nematodes live in fresh and brackish
waters, and transitional zones. Many of these species
tolerate rapid fluctuations in salinity. As in marine habitats,
the animals are usually present in the sediment, although
they may occasionally swim freely. The most dense
nematode faunas are associated with a reasonable oxygen
supply and sediment with a high organic content. Lakes
have a very variable fauna which probably depends upon
their physical attributes, such as isolation and thermal
stratification. Shallow marginal waters may be quite rich,
probably sharing some species with wet terrestrial habitats.
However, deeper waters seem to be species-poor unlike
marine systems. A notable exception to this is Lake Baikal,
where, as among other animal groups, considerable
speciation has occurred and endemism appears to be high
(Nicholas, 1984).
In the soil the distinction between microbotrophic and
parasitic nematodes becomes very blurred in certain taxa.
All kinds of soils support large nematode communities (see
Table 9.2 below) and the richest tend to be where there is
plenty of organic matter, fine plant roots, etc. The
interactions with plant roots and other organisms, such as
fungi, are extremely complex and difficult to assess.
It is thought that parasitism has arisen independently in
several nematode taxa, and certainly the microbotrophic
forms illustrate a great variety of interactions which could
Nematodes
be considered as stages in the evolution of parasitism. For
example, there are many examples of phoretic relationships
with invertebrates. These range from larval stages attaching
externally to mobile hosts who carry them to the next food
source, to larval stages which live within a host apparently
without harming it, but which cannot escape to continue
their life cycle until the host dies of natural causes. Many
of these relationships are very finely tuned to the life cycle
of a specific carrier whilst others use a variety of suitable
invertebrates. Not all relationships benefit the nematode
alone: in some cases the carrier may also feed upon the
nematodes. Relationships with plants may be equally
complex, as nematodes may often feed upon the bacteria on
and in decaying roots. However, some species are suspected
of spreading disease to increase their food resource or of
being able to feed upon living plant tissue as an alternative
to bacteria. Even within one species, different forms may
show different degrees of interaction, making rigid
definitions impossible.
Predaceous nematodes
Predaceous nematodes are found in all habitats but are most
abundant in terrestrial systems. All eat a few to many
multicellular organisms in the course of their development,
although bacteria, ciliates and organic particles may also be
eaten. Little is known about prey-specificity in nature, as
most studies, by necessity, have been carried out under
laboratory conditions. However, some extremely common
groups include other nematodes as prey items and may be
potential biological control agents for nematode pests of
plants. For instance, a single nematode of the family
Mononchidae has been observed to kill over 1,000
nematodes in a three-month period and estimates of density
suggest that up to 300 million mononchid nematodes might
be contained in an acre of soil (Poinar, 1983). However,
observations also suggest that almost any invertebrate of the
correct size may be eaten and prey location is a chance
affair.
Little is known of the aquatic predaceous nematodes.
However, observations which suggest that some marine
forms may be able to penetrate foraminiferan tests to get at
the body inside are of considerable interest as borings
similar to those attributed to these nematodes have been
seen in fossilized foraminiferan tests from the Holocene and
Cretaceous periods (Poinar, 1983).
Table 9.2 Abundance and biomass of soil nematode fauna from different types of
ecosystem
ECOSYSTEM ABUNDANCE x 1000m? BIOMASS*
MEAN RANGE MEAN RANGE
Tundra 3,490 800-10,000 1,350 265-4,130
Coniferous forest 3,330 1,125-15,000 510 180-1,696
Eucalyptus forest 5,467 4,040-7,449 1,423 770-2,050
Deciduous forest 6,270 255-29,800 2,760 75-15,200
Temperate grassland 9,190 2,432-30,000 3,800 650-17,800
Fen, bog, heathland 1,660 330-3,900 660 350-900
Desert 760 423-1,100 410 125-700
Tropical forest 1,700 1,500-1,900 - -
Source: Sohlenius, B. 1980. Abundance, biomass and contribution to energy flow by soil nematodes in terrestrial ecosystems. Oikos 34:186-94.
Note: * biomass is measured here in mg weight per m?.
89
1. Biological Diversity
Table 9.3 Distribution of nematode genera among groups of vertebrates
FISH AMPHIBIANS REPTILES BIRDS MAMMALS HOST SPECIES
PER NEMATODE
GENUS
Fish 62 6 7 4 1 250
Amphibians 20 22 2 1 50
Reptiles 62 4 4 60
Birds 113 17 60
Mammals 387 8
TOTAL GENERA 80 51 99 140 410
Source: Modified from Inglis, W.G. 1965. Patterns of evolution in parasitic nematodes. In: Taylor, A.E.R. (Ed.), Evolution of Parasites. Blackwell
Scientific Publishers, Oxford, UK and Poinar, G.O. 1983. The Natural History of Nematodes. Prentice-Hall Inc., New Jersey, USA.
Note: Numbers underlined indicate genera exclusive to each vertebrate group.
Parasitic nematodes
Plant parasitic nematodes have been found in most species
of terrestrial plants, all over the world. Many are
polyphagous and consequently a plant species may be
attacked by a wide range of nematode species. For instance,
Poinar (1983) lists 36 nematode species in 15 genera which
have been identified parasitising potatoes and six species in
four genera from wild chicory. Most fungi also suffer from
nematode attacks, some species being serious pests in
mushroom culturing operations. Plant parasites are
apparently much less common in aquatic habitats, and
Telatively few species are known from seaweeds and marine
fungi.
Parasitic nematodes are similarly widespread in both
invertebrate and vertebrate hosts and have evolved some
remarkably complex life cycles. The greatest number of
invertebrate parasites known are in the insects and some of
these have been studied in great depth in the hope of
developing successful biological control methods. As with
the plant parasites many nematode species can attack a wide
range of insect hosts. Others are highly specialised and
adapted to the life cycle of one particular host. Of the
former group, two nematode families include genera which
have evolved mutualistic relationships with a single
bacterium genus, which is unkown in a free living state.
These nematodes introduce bacterial cells into a host insect
which dies soon after becoming infected. The bacteria then
grow on the body and the nematodes feed on the bacteria,
ensuring some are carried to infect a new host. Insects of
Table 9.4
CENTRAL AMERICA SOUTH AMERICA BRAZIL
AND CARIBBEAN
% % %
Crop loss Crop loss Crop
Tomato 38 Cucumber 33 Tomato
Chayote 38 Tomato 27 Coffee
Guava 35 Bean (common) 24 Soybean
Pumpkin 22 Watermelon 23 Cotton
Bean (common) 16 Pepper 22 Papaya
Yam 16 Eggplant 20 Yam
Mean % loss
(all crops) 15 15
ten different orders are known to be attacked by these
species (Poinar, 1983).
Vertebrate parasites are equally widespread and, here again,
some may utilise a whole range of hosts whilst others are
extremely host specific. Many have developed complicated
methods of dispersal which may involve invertebrates (or
occasionally other vertebrates) as intermediate hosts. Some
of the world’s most debilitating diseases are spread by this
method, such as onchocerciasis (river blindness).
Nematode parasites tend to become more specialised in
more developed vertebrate groups. The majority of
nematode genera are confined to a single vertebrate genus.
Of those that do have a wider host range a few can utilise
different classes, but most are restricted to similar animals.
Table 9.3 illustrates the higher diversification of nematode
parasites in the higher vertebrate groups and the greater
specifity associated with this.
THE ECOLOGICAL
NEMATODES
IMPORTANCE OF
Free living nematodes are vital components of ecosystems.
Although not themselves decomposers, many feed on the
primary decomposers, the bacteria and fungi, which break
down complex organic molecules and thus make these
nutrients available in the food chain again. They are
therefore elementary in the decomposition cycle. The
predaceous species are also important consumers near the
Estimated crop losses due to Meloidogyne species in tropical regions
WEST AFRICA SOUTHEAST ASIA
% %
loss Crop loss Crop loss
25 Tomato 46 Tomato 24
24 Cowpea 43 Melon 18
23 Okra 42 Bean (common) 18
17 Carrot 38 Eggplant 17
15 Pigeon pea 35 Black pepper 16
15 Melon 33 Chinese Pechay 16
13 25 11
Source: Adapted from Sasser, J.N. 1979. Economic importance of Meloidogyne in tropical countries. In: Lamberti, F. and Taylor, C.E. (Eds),
Root-knot Nematodes (Meloidogyne species). Academic Press, London, UK.
Note: Only the six worst-affected crops are shown in each case.
90
base of food webs, feeding on unicellular algal primary
producers and smaller metazoans.
Nematodes are most often studied in their destructive
capacity, as pests of agricultural crops and as parasites of
livestock and humans. However, there is also potential for
biological control applications, against a wide range of
insect pests and against other nematode species. Free-living
nematodes have been used as models for various
experiments on the functioning of ecosystems and as
indicators of environmental health, such as water pollution.
Other potential and actual uses include nematodes as
indicators of the quality of terrestrial soils, freshwater and
marine sediments (van der Wal and de Goede, 1988), and
in a whole range of biological research projects (see
Nicholas, 1984, for examples).
There are many aspects to the problems of nematode
association with crops. For instance, nematode species
which are useful in controlling pathogenic root fungi in one
situation may in another destroy mycorrhizal fungi,
necessary for good plant growth. Similarly, some
nematodes which feed harmlessly or even usefully on
bacteria most of the time may also be able to move into
plant roots, either to eat healthy plant tissue directly or to
infect them to provide more food for their bacteria. This
change may depend on environmental conditions, for
instance the soil drying out, and may produce a sudden
reaction in the crop which superficially resembles water
stress. These cases are the cause of some debate and
considerable research. However, other nematodes are
without doubt serious crop destroyers. Poinar (1983) quotes
estimates which suggest that 7-15% of the annual crop
production of the USA is destroyed by nematodes. Table
9.4 shows the estimated yield losses in several tropical
regions due to species Meloidogyne, one of the most
destructive nematode genera. Only the six worst-affected
crops in each region are shown here in detail but in the
original table Sasser (1979) gives figures for up to 21 crops
Nematodes
in each region. The most destructive species in each case is
M. incognita, followed by M. javanica, M. arenaria and M.
hapla.
Some of these problems have arisen as a result of crop
monoculture which reduces natural control systems that
normally keep such pests within acceptable limits. Various
methods of control are possible, including timed planting to
miss the most active cycle of the parasite, crop rotations
which can include crops poisonous to the nematodes, and
flooding. Another form of natural control which has
received considerable attention in recent years entails use of
fungi that are predaceous or parasitic upon nematodes. At
least one of these former, a trap-forming deuteromycete in
the genus Arthrobotrys, is commercially available (Poinar,
1983) and is effective in tomato fields and greenhouses
against Meloidogyne species. Various other fungi have been
tested with varying results and other fungi which apparently
produce nemotoxins are also being studied.
On the other hand, control by nematodes of fungal plant
diseases and weeds have been investigated. For example, an
encysting plant parasite Paranguina picridis has been used
with some success in the USSR to control knapweed
(Poinar, 1983). Predaceous nematodes have also been
considered as control agents for ectotrophic root parasites,
especially other nematodes, and microbotrophic nematodes
for control against certain infective bacteria.
In contrast to plant parasites, the invertebrate parasites are
rarely a problem to man (except where plants or higher
animals are also part of the life-cycle). In fact many have
great potential for control of pest insects. In particular,
certain nematodes have been intensively studied for possible
mosquito control and others which parasitise water snails
may be able to control schistosome-bearing snails. Insect
pests of crops and livestock are also targeted by research
programmes; several examples which have been tried are
shown in Table 9.5, adapted from Poinar, 1983.
Table 9.5 Examples of nematode species investigated as biological control agents
FAMILY SPECIES INSECT PEST LOCATION HABITAT
Mermithidae Romanomermis mosquitoes North America, Taiwan, Europe, Ponds,
culicivorax Africa, Oceania, Central ditches,
America, Thailand lakes.
Diplogasteridae Pristionchus Colorado beetle Poland Soil
uniformis
Steinernematidae Neoaplectana Japanese beetle Eastern USA Soil
glaseri
Heterorhabditidae Heterorhabditis Agriotes spp. Italy Soil
bacteriophora (click beetles)
Neotylenchidae Deladenus Sirex noctilio Australia Trees
Siricidicola (wood wasp)
Allantonematidae Heterotylenchus Musca autumnalis North America Dung
autumnalis (face fly)
Sphaerulariidae Tripius sciarae Sciarid flies England (greenhouse) Soil
Source: Adapted from Poinar, G.O. 1983. The Natural History of Nematodes. Prentice-Hall Inc., New Jersey, USA.
1. Biological Diversity
Table 9.6 Estimates of nematode infections in man (in millions)
DISEASE NEMATODE/S AFRICA ASIA CENTRAL OCEANIA NORTH EUROPE ‘USSR’
(excl. & SOUTH AMERICA (excl.
‘USSR’) AMERICA ‘USSR’)
Ascariasis Ascaris lumbricoides 159 931 104 1 5 39 30
Hookworms (various) 132 685 104 2 3 2 4
Human Enterobius vermicularis 24 136 40 1 29 75 48
pinworm
Trichuriasis Trichuris trichiura 76 433 94 1 1 41 41
Trichinosis Trichinella spiralis 1 3 35 5 2
Others 9 49 21 <1 1 1 3
Elephantiasis Wuchereria bancrofti and 59 300 22 2
Brugia malayi
Other filariae 178 57 39
Source: Peters, W. 1978. Comments and discussion Il. In: Taylor, A.E.R. and Muller, R. (Eds), The Relevance of Parasitology to Human Welfare
Today. Blackwell Scientific Publications, Oxford, UK.
Nematode parasites of vertebrates are an enormous drain
upon human resources, both in the effects on domestic
animal species and on human life directly. The World
Health Organization produces estimates for the numbers of
people afflicted with the major parasitic diseases. Poinar
(1983) gives figures for four of these for 1977-78:
hookworm disease, onchocerciasis, ascariasis and
trichuriasis in Africa, Asia and Latin America. Of these the
first two cause the greatest number of deaths each year: 50-
60 thousand deaths among 7 million to 900 million people
with hookworm disease, and 20-50 thousand deaths out of
30 million estimated cases of onchocerciasis. A different
presentation of similar data is given in Table 9.6, adapted
from Peters (1978). These estimates are apparently based on
data collected in the 1940s although Peters suggests they
adequately represent the current situation.
The monetary costs caused by livestock disease are also
immense, in terms of prevention, treatment, animals lost
and human time. Where these parasites are also
transmittable to humans, such as several of those affecting
pigs, precautions against infection are also costly and time-
consuming. Thus, unlike the possible benefits from free
living and plant parasitic nematodes, and the considerable
potential in invertebrate parasites, there are no obvious uses
92
of vertebrate parasites with benefit to humans.
References
Anderson, R.V. 1984. The origins of zooparasitic nematodes.
Canadian Journal of Zoology 62:317-28.
Barnes, R.D. 1980. Invertebrate Zoology, 4th edn. Holt-Saunders
Tokyo, Japan. 1,089pp.
Inglis, W.G. 1965. Patterns of evolution in parasitic nematodes. In:
Taylor, A.E.R. (Ed.), Evolution of Parasites. Blackwell Scientific
Publishers, Oxford, UK. Pp.79-124.
Nicholas, W.L. 1984. The Biology of Free-living Nematodes, 2nd edn.
Clarendon Press, Oxford, UK. 251pp.
Peters, W. 1978. Comments and discussion I]. In: Taylor, A.E.R. and
Muller, R. (Eds), The Relevance of Parasitology to Human Welfare
Today. Blackwell Scientific Publications, Oxford, UK. Pp.25-40.
Poinar, G.O. 1983. The Natural History of Nematodes. Prentice-Hall
Inc., New Jersey, USA. 323pp.
Sasser, J.N. 1979. Economic importance of Meloidogyne in tropical
countries. In: Lamberti, F. and Taylor, C.E. (Eds), Root-knot
Nematodes (Meloidogyne species). Academic Press, London, UK.
Pp.359-374.
Sohlenius, B. 1980. Abundance, biomass and contribution to energy
flow by soil nematodes in terrestrial ecosystems. Oikos 34:186-94.
Wal, A.F. van der and Goede, R.G.M. de (Eds) 1988. Nematodes in
Natural Systems. Report of a workshop held at the Dept. of
Nematology, Agricultural University, Wageningen, The
Netherlands, 16-18 December 1987. Mededeling 199.
10. DEEP-SEA INVERTEBRATES
DEEP-SEA COMMUNITIES
Until the mid-1960s it was believed that oceanic diversity
was concentrated in shallow water around coasts and
declined with both depth and distance from land as food
resources became more remote. The first reports of
unexpectedly high species diversity in bottom living
communities arrived in 1967 with samples collected using
a new technique: the epibenthic sled (Hessler and Sanders,
1967). Although many were initially sceptical of the
conclusions, the deep-sea environment has been an active
area of research and is now known to support communities
rich in species, high in endemism and often ecologically
unique. In terms of species numbers alone, the marine
environment provides a relatively minor proportion of the
global total.
Approximately 71% of the Earth’s surface is covered by
sea, and about 51% of its surface by ocean over 3,000m in
depth. Deep-sea communities are thus prevalent over a
major proportion of the planet. All deep-sea habitat is in the
aphotic zone, well below the distance sunlight can
penetrate. Community structures and food webs are
therefore very different from those found on land and in the
shallower parts of seas in that, except in the specialist case
of hydrothermal vents (described below), there is no
primary production and all life relies on organic material
from other parts of the ocean. As deeper and deeper levels
are reached biomass falls exponentially (Rowe, 1983). This
was misinterpreted as being synonymous with falling
species diversity (Grassle, 1991). Because, despite their
enormous volume, the deep oceans appear to be relatively
simple ecosystems, there was little reason to imagine that
they should make any significant contribution to overall
global species diversity. That species diversity in the
benthic community should rise with increasing depth was
therefore a major discovery.
The benthic samples taken by Hessler and Sanders (1967)
and later workers have revealed a hitherto unexpectedly
high species richness. This discovery has prompted
speculation that the deep sea is a site of prolific speciation
and, as one of the most stable and ancient environments on
Earth, perhaps the origin of certain higher-level taxa (Gage
and Tyler, 1991). Several ideas have been postulated to
explain this high diversity but it would appear that a
combination of factors is important. Grassle (1991) suggests
four major influences:
© the relative lack of environmental extremes such as those
of temperature, salinity, low oxygen and major
disturbances
© patchy food resources
e local disturbances and structures caused by animal
activities
© a large area with few barriers to dispersal.
The first three of these are equivalent to the processes
thought by some to be fundamental to the high species
diversity in tropical terrestrial and shallow water
ecosystems. Environmental stability allows the development
of high species diversity with many highly specialised
93
Deep-Sea Invertebrates
species. This is supported by observations in deep-sea areas
that do not have long-term environmental stability, such as
trenches and areas of strong bottom currents; these usually
have a much reduced species diversity although their faunas
may be of interest in other ways (Thorne-Miller and
Catena, 1991).
The patchiness of food availability and local disturbance can
be compared to the importance of gap appearances in the
canopy of tropical forests, both involving small scale habitat
diversity within a larger homogenous area and the
maintenance of a mosaic of disequilibrium populations
(Grassle, 1989). Most organisms which live in the deep sea
are totally dependent on organic detritus falling from
euphotic zones. This is largely of planktonic and faecal
origin but larger masses such as pieces of wood, carcasses
and algal mats are also of importance. Local disturbances
such as feeding activities and burrowing and mound-
building by polychaete worms also ensure local topographic
variations which provide a variety of microhabitats. Weak
bottom currents allow particulate organic matter to
concentrate in hollows and lees.
The large area of the deep ocean zone, coupled with the
above factors, results in a very large species pool with wide
dispersion potential. Grassle (1991) estimates that if the
currently observed species-area relationship is extrapolated
the total species pool may be in the order of 10 million.
Although, as discussed below, there are many problems
with predictions of this type, even this figure may be
conservative.
Faunal composition
Studies of the benthic species assemblages of different
regions are still in their infancy. The major difficulty is
obtaining quantitative samples, since the depths involved are
far greater than a diver can go. Much of the work which
has been carried out has not been coordinated, leading to
different sieve sizes for sampling, different collection
techniques and different assessments of biomass (Rowe,
1983). This makes comparisons between sites difficult. In
addition, taxonomic problems in certain taxa have meant
that, while it may be possible to have a species count from
any one sample, it is not possible to say what the similarity
is between samples. Nearly half the species in each new
sample may be undescribed (Grassle, 1989), and there may
be few taxonomists working on any one group, raising
problems of species identification. Even in well sampled
areas, sample sizes are small compared to the regions they
are supposed to represent, and it is uncertain to what extent
results can be extrapolated. However, the rate of discovery
of new species and the proportion of species currently
known from only one sample both indicate that a great
number remain to be discovered (Grassle, 1991).
Benthic fauna is usually classified into size classes,
increasing from the nanobiota, through the meiofauna, the
macrofauna and finally to the megafauna. A problem with
this type of classification is it splits natural taxonomic
groups and even age classes of the same species. Many
workers prefer to classify all the members of certain taxa
1. Biological Diversity
into the size class which best represents the group; for
example, all nematodes are often considered as meiofauna.
Small size and taxonomic problems mean that few
comparative data are currently available on meiofaunal
diversity. The major taxonomic groups in this size class are
the nematodes and foraminiferans (protozoa). Other
important taxa in this size class include the harpacticoid
copepods and ostracods.
More information is available for the macrofauna, which
has been more extensively studied than other size classes.
This is typically dominated by polychaetes (up to 75%
numerically), peracarid crustaceans (including cumaceans,
tanaids, isopods and amphipods) and a variety of smaller
molluscs (Gage and Tyler, 1991), but most other phyla are
also represented. Figures from Grassle (1991) (see Table
10.1 and Fig. 10.1) demonstrate the species, family and
phylum composition of a typical sample; (however, note
these samples were taken at bathyal rather than abyssal
depths - see below). Wolff (1977) also provides examples
of the taxonomic composition of the macrofauna (and
megafauna using the taxa listed here) in a number of
regions (Table 10.2 and Fig. 10.2). However, the sampling
methods are not the same in each area so these results may
not be comparable.
Table 10.1 Diversity in benthic
samples *
NO. OF NO. OF
GROUP FAMILIES SPECIES
Annelida 49 385
Arthropoda 40 185
Mollusca 43 106
Echinodermata 13 39
Nemertina 1 22
Cnidaria 10 19
Sipuncula 3 15
Pogonophora 5 13
Hemichordata 1 4
Echiura 2 4
Priapulida 1 2
Brachiopoda 1 2
Ectoprocta 1 1
Chordata 1 1
TOTAL 171 798
Source: After Grassle, J.F. 1991. Deep-sea benthic biodiversity.
Bioscience 41(7).
Note: * Sea-bed samples from 1,500m to 2,500m depth off New
Jersey, north-east Atlantic.
Table 10.2 Composition of benthic macrofauna (percentage of total species present)
LOCATION TRENCHES* CENT.N PACIFIC NW ATLANTIC NW ATLANTIC
SAMPLE TYPE TRAWL A.D. A.D. E.S.
DEPTHS (m) 6,000-10,000 5,600 4,400-5,000 4,700
Polychaetes 7; 55 55 8
Peracarid crustacea (total) 5 24 33 32
Tanaidacea <1 18 19 1
lsopoda 4 6 12 18
Amphipoda <1 (e) 2 5
Bivalvia 19 7 4 47
Echinodermata (total) 57 1 1 2
Ophiuroidea 2 <1 <1 72
Holothuroidea 54 <1 (0) {e)
Others 11 11 8 11
Number of individuals 21,589 287 681 3,737
Source: After Wolff, T. 1977. Diversity and faunal composition of the deep-sea benthos. Nature 267:780-785.
Note: * Trenches = Kurile-Kamchatka, Japan, Kermadec and Java A.D. = Anchor Dredge; E.S. = Epibenthic Sledge.
In the megafauna, echinoderms of several classes are often
the dominant mobile (or errant) life forms on or in
association with the sea bottom. Their distribution may be
very uneven, they may sometimes occur in great numbers
on patches of detritus fallout and some scavenging forms
may be found in large congregations at bait. Giant
scavenging amphipods, growing up to about 18cm in length,
are also characteristic in many areas. However, the high
mobility of these animals means they are rarely caught in
trawls and have been less well studied than less active
animals. Other arthropods include a variety of sea spiders
(Pycnogonida) and decapods of several families (both errant
and sessile). Errant animals of several other taxa occur,
including polychaetes, hemichordates, cephalopods and fish.
Sessile animals generally occur on any suitable surfaces.
Sponges (Porifera), especially the glass sponges, are widely
distributed and coelenterates (Cnidaria) are also well
94
represented, dominated by anthozoans. Other taxa include
bryozoa and brachyopoda.
Distribution of deep ocean biodiversity
There are general trends in species richness with respect to
depth in benthic communities. The picture is very
incomplete, however, so conclusions must be tentative. Rex
(1983) examined data from four major taxonomic groups
(polychaetes, gastropods, protobranchs and cumaceans)
along a depth gradient down to 5,000m. All showed
maximum diversity between 2,000m and 3,000m. The three
of these taxa which had been sampled with an epibenthic
sled rather than an anchor dredge also showed a higher
diversity between 4,000m and 5,000m than between 0m and
1,000m (polychaetes were the exception). However,
whether this is an artefact of the difference in sampling
technique or a true difference between the taxa is unclear.
The assemblages of different depth zones have differing
patterns of geographical distributions. Abyssal species
appear to have the most widespread distributions (Angel,
1991), probably because there are fewest barriers to larval
dispersal. For example, in the Polychaeta, which is one of
the less cosmopolitan groups, 78% of all North Atlantic
abyssal species are found in both the East and West
Atlantic, compared to 58% of the bathyal species (Gage and
Tyler 1991). Hadal, or ultra-abyssal, communities again
have more disjointed distributions as only about 1% of the
Earth’s surface is covered with water of such depth. Plain
communities at these depths are poorly sampled but of
particular interest are trench faunas (although not all
trenches reach hadal depths). These are dealt with
separately below.
Latitudinal patterns are even less well studied. In pelagic
communities there is a general trend for the number of
species to increase from the polar to the tropical regions.
Buzas and Culver (1991), also report a definite latitudinal
gradient in the foraminiferans of open ocean sediments,
typically ranging from 10-30 species in a few millimetres of
sediment at high latitudes to 50-70 species in tropical
latitudes. Whether this pattern is representative of the
benthos as a whole is unclear.
OCEAN TRENCHES
Physical evolution and properties
Ocean trenches are formed as a consequence of plate
tectonic processes where sectors of expanding ocean floor
Deep-Sea Invertebrates
pushes upon an unyielding continental mass or island arc,
resulting in the crust buckling downwards (subducting) and
being destroyed within the hot interior of the Earth. As
oceanic crust ages and cools, it becomes denser and stiffer,
resulting in a steeper angle of subduction and a deepening
trench. Fig. 10.3 shows the locations of the principal
known trenches, those occurring along the western edge of
the Pacific being both the deepest, and geologically the
oldest. Seismically, ocean trenches are highly active, as
subduction is an erratic rather than a smooth process. This
results in an unstable and unpredictable habitat compared to
the relative environmental stability of the adjacent abyssal
plains (Angel, 1982).
Being generally close to land masses, ocean trenches tend
to have relatively high rates of sedimentation, a significant
amount of which is of organic origin and an important
available food source for trench communities. Several
trenches also underlie highly productive cold water
upwelling zones, the organic fallout from which contributes
greatly to their richness. The water within trenches
generally originates from the surrounding bottom water,
which is derived from cold surface water at high polar
latitudes and is relatively well oxygenated (Angel, 1982).
Endemism, diversity and biomass
Trenches tend to be isolated linear systems. This, combined
with their high seismic activity, would suggest that faunas
low in species diversity but relatively high in numbers of
endemic species should be found. These would be expected
to show strong affinities at generic and family levels to
other trenches in the same system, having all originated
from the same parental species inhabiting the surrounding
Table 10.3. Endemism among hadal species
GROUP TOTAL NO. OF NO. OF SPP. % ENDEMIC
HADAL SPECIES EXCLUSIVELY AT HADAL SPECIES
>6000m DEPTHS >6000m
Cumacea 3 3 100.0
Harpacticoida 2 2 100.0
Ostracoda 2 2 100.0
Crinoidea 9 8 88.9
Gastropoda 16 14 87.5
Pogonophora 26 22 84.6
Amphipoda 17 14 82.4
Tanaidacea 19 15 78.9
lsopoda 49 37 75.5
Porifera 12 <) 75.0
Coelenterata Uz 9 75.0
Pisces 4 3 75.0
Bivalvia 26 17 65.4
Holothurioidea 22 14 63.6
Echiurida 8 5 62.5
Ophiuroidea 5 3 60.0
Asteroidea 12 6 50.0
Others 5 2 40.0
Polychaeta 32 12 37.5
Cirripedia < 1 33.3
Pycnogonida 3 1 33.3
Foraminifera 126 35 27.8
Sipunculida 4 {e) 0.0
Total spp. 417 234 56.1
Excl. Foraminifera 291 199 68.4
Source: Wolff, T. 1970. The concept of the hadal or ultra-abyssal fauna. Deep-Sea Research 17:983-1003.
1. Biological Diversity
Figure 10.1 Species and family diversity in sea-bottom samples
Number of species/families
Species | Families
Ls] Ls} Lt o oO i) 9 o A Le] ie) oO fs) oO
1) UD oO ~ c = = i L a 50) D y -
= 2° wo Q = L 3 ce) 2 Ls} ° = oO Lo
= Q 2 v SG Qo £ 2 Be) a os ° v
a ° = L e me] = Q = i ° =) L L
ic ‘ = o g = =) fo} 0 5 = a Q fe}
c <£ g UD S a i= w £ £ o ° =
< + fe} 9 ro) = fe) y) s) = Pe) re)
L c 72) fo = o & 5)
=< = fe) [= L w
£ a ¥ o
i} Phyla
Source: Grassle, J.F. 1991. Deep-sea benthic biodiversity. Bioscience 41(7).
Note: Samples taken at 1,500-2,500m depth off New Jersey, USA.
Figure 10.2 Composition of benthic macrofauna
Percentage of total number of individuals
Trenches CTrawl) Cent .N.Pacific CA.0.) N.W.Atlantic CA.0.) N.W.Atlantic CE.S.)
6,000-10,000 m 5.600 m 4,400-S,000 m
BEE Peracar id
Polychaetes Sache Bivalves
“te crustaceans
Source: Wolff, T. 1977. Diversity and faunal composition of the deep-sea benthos. Nature 267:780-785.
96
A
Echinoderms ca Others 2
= 5
.700 m
Anchor dredge
Epibenthic siedge
Deep-Sea Invertebrates
Figure 10.3 Distribution of the main ocean trenches
(2861 ‘jeBuy woy payipow)
YoIMpues UNoS
C
SepuqeH MeN
f ay eugueweig
Sat
p
depeuLiey / a _
allu9-Tua,
AC essa geen ( : ‘\
ener
=e 2 ulewg mon ——> x =<
eyouewioy Rt wi deh pe SD ip \ sofeu9
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URDUOWY e1PPIN eve
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— =
1. Biological Diversity
abyssal plains (Angel, 1982). In the few trenches studied,
these hypotheses would appear to be true; however, as with
other aspects of deep-sea diversity, generalisations remain
tentative.
Angel (1982) quotes Professor G.M. Belyaev (from which
the following paragraph of information is taken). Between
50% and 90% of the fauna of each ocean trench is endemic,
compared to the overall endemism of hadal, or ultra-abyssal
faunas, which is in the region of 57-60% (for example, see
data given by Wolff, 1970, Table 10.3). There are some 25
known endemic hadal genera, representing some 10-25% of
the total number of genera occurring in the hadal zone, and
two known endemic hadal families; the Galatheanthemidae
(Actinaria) and Gigantapseudidae (Crustacea). The latter
family contains a single species: Gigantapseudes adactylus.
The greatest number of endemic species known from a
single trench is a sample of 200 from the Kurile-Kamchatka
Trench; this may be compared with 10 endemic species
known from the Ryukyu and Marianas Trenches. The
Banda Trench has the lowest recorded proportion of species
endemism (33%), and is probably the youngest trench
geologically. In total, representatives of 33 classes, 150
families and about 240 genera are known from hadal
depths.
As noted, high seismic activity may tend to produce low
species diversity. Rapid sedimentation may have a similar
effect. For example, the Aleutian Trench and the Japan
Trench have relatively low macrofaunal diversities,
attributable to frequent catastrophic slumping of canyon
wall sediment (Grassle, 1989).
In general, comparative data are sparse because of the
variety of collection techniques employed. The composition
of trench faunas is unusual (compared to abyssal faunas) in
that they tend to be dominated by deposit-feeders (Angel,
1982) and show a higher percentage of species of
amphipods, polychaetes, bivalves, echiurids and
holothurians, and a lower percentage of sea _ stars,
echinoids, sipunculids and brittle-stars, and especially non-
actinian and scyphozoan coelenterates, bryozoans,
cumaceans and fishes, than in the surrounding abyss.
Decapod crustaceans are completely absent (Gage and
Tyler, 1991).
Trenches appear to have a higher biomass than adjacent
shallower areas, although within the trenches themselves the
stocks of macrofauna decrease with depth at a rate similar
to the general declining pattern. The higher biomass in
trenches is probably a reflection of the net accumulation of
sediment from the adjacent shallow continental margins
(Rowe, 1983), as the amounts of available nutrients have a
profound effect on trench faunas; 8.8g/m? of living
organisms have been assessed from the nutrient-rich South
Sandwich Trench and 3.44g/m? from the Kurile-Kamchatka
Trench, compared to 0.008g/m? from the nutrient-poor
Marianas and Tonga Trenches (Angel, 1982).
HYDROTHERMAL VENTS
Hydrothermal vent communities were first discovered in
1977, at a depth of 2,500m on the Galapagos Rift. They are
now known to be associated with almost all known areas of
98
tectonic activity at various depths (see Fig. 10.4). These
include: along the East Pacific Rise off Mexico, in the
Guaymas Basin in the Gulf of California, on the Juan de
Fuca Ridge off Washington State, in subduction areas off
Oregon and Japan, on the Mid-Atlantic Ridge at 26°N, in
the Mariana Trough near the Mariana Trench, and in the
Lau and North Fiji Basins to the west and east of Fiji (Gage
and Tyler, 1991). These tectonic regions include ocean-
floor spreading centres, subduction and fracture zones, and
back-arc basins (Gage and Tyler, 1991). Cold bottom-water
permeates through fissures in the ocean floor close to
ocean-floor spreading centres, becomes heated at great
depths in the Earth’s crust and finds its way back to the
surface through hydrothermal vents. The temperature of
vent water varies greatly, from around 23°C in the
Galapagos vents, to around 350°C in the vents of the East
Pacific Rise, and they may be rich in metalliferous brines
and sulphide ions (Angel, 1982). Although the vent water
may be at a high temperature, the majority of species live
out of the main flow at temperatures of around 2°C, the
ambient temperature of deep-sea water.
Although vent communities are often separated from one
another by gaps of a kilometre or so, they can be up to
100km apart. They have yet to be found in certain areas of
known hydrothermal activity, such as the Red Sea (Grassle,
1986). Hydrothermal vents and their associated communities
are relatively short-lived at any particular site, probably
only being active for between several years and several
decades. This has been suggested by discoveries of ‘dead’
vents (visible from the remains of white shells which
dissolve away completely in about 15 years) and by growth
measurements of individual organisms (indicating very rapid
growth to maturity at a large size) (Gage and Tyler, 1991).
However, active hydrothermal centres appear to move
relatively slowly, thus allowing dispersal of vent organisms.
Areas of tectonic activity are connected over most of the
earth’s surface, and although this network is in a dynamic
state, new areas are linked to old and so vent communities
could be part of a unique ecosystem at least 200 million
years old (Grassle, 1985). Studies on variation in vent
species, comparing those in the main network and those
isolated in remote parts of the system, provide important
opportunities for evolutionary and genetic studies. Vent
species are also of interest in that they flourish in the dark
at high pressures and low temperatures (Grassle, 1986),
which previously had been thought to inhibit productivity.
Hydrothermal vent communities are unique in that they are
supported by a non-photosynthetic source of organic carbon,
i.e. chemosynthetic primary production. The enriched
hydrothermal fluid supports large numbers of bacteria
(predominantly Thiomicrospira species) which form dense
bacterial ‘mats’, and are capable of deriving energy from
reduced compounds such as hydrogen sulphide (Grassle,
1986, Gage and Tyler, 1991). Many of the vent species
filter-feed on these bacteria, whilst others rely on symbiotic
sulphur bacteria for energy (Angel, 1982).
Endemism, diversity and biomass
The overall species diversity at vents is low compared with
other deep-sea soft-sediment areas (Grassle, 1986), but
endemism is high. More than 20 new families or sub-
Deep-Sea Invertebrates
Figure 10.4 Hydrothermal vent and cold seep communities
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99
1. Biological Diversity
families, SO new genera and nearly 160 new species have
been recorded from vent environments, including brine and
cold seep communities (discussed below) (Grassle, 1989;
Gage and Tyler, 1991). Examples of these new taxa are
given in Table 10.4.
In biogeographic terms, vents can be regarded as
ephemeral, biogeographic islands. With increasing spatial
separation the species composition can vary considerably,
with some species being replaced by closely related forms.
Differences in subsurface flux of hydrothermal fluids and in
vent configuration can result in large differences in faunal
composition over short distances within or between vent
fields. Geochemical differences between vent communities
may also result in faunal dissimilarities, between the
Galapagos and East Pacific Rise vents, for example
(Grassle, 1986). However, the major features of the fauna
at each vent site are consistent, whilst none of the species
seems to be ubiquitous. The larvae of many vent species
appear to have relatively poor dispersal abilities (Grassle,
1986), and this could contribute to maintenance of high
endemism.
The biomass of vent communities is usually high compared
to other areas of similar depth, and varies according to
water temperatures and chemistry, reaching 8.5kg wet
weight per m? at lower temperature vents, and averaging 2-
4kg wet weight per m? at the hottest vents (200-360°C)
(Gage and Tyler, 1991). Dense colonies of tube-worms,
clams, mussels and limpets typically constitute the major
proportions of biomass. Swarms of the probably vent-
specific copepod species /saacsicalanus paucisetus reached
densities of 920 individuals m ? and a dry weight biomass
of 133 mg? at one site. Microbial production at low-
temperature vents (10°C) is thought to be two or three
times that of photosynthetic production at the surface in the
same region (Gage and Tyler, 1991).
Features of some major vent regions are noted below.
Galapagos Spreading Centre
This consists of 12 known active populated vents and three
‘dead’ vents along a 30km section of ridge-crest. The two
large bivalves Calyptogena magnifica and Bathymodiolus
thermophilus, and vestimentiferan worms (especially the
tube-dwelling Riftia pachyptila) are the most distinctive
species of these hydrothermal vents (Grassle, 1986).
Eastern Pacific Rise
These hydrothermal vents support a similar fauna to the
Galapagos Spreading Centre, including the same two
bivalve species (which can occur in enormous densities - the
biomass of B. thermophilus may exceed 10kg/m?), and
Riftia pachyptila. More than 30 species of limpet-like
gastropod have been recorded (mostly as yet undescribed),
and mussels, shrimp, anemone and limpet species (Gage
and Tyler, 1991). The spreading rate of 11-12cm/year is
greater than that of the Galapagos spreading centre
(Grassle, 1986).
Mid-Atlantic Ridge
The active hydrothermal vents discovered on this ridge are
characterised by the presence of two species of caridean
shrimp belonging to the new family Bresiliidae. These
100
occur in great numbers, along with mats of bacteria.
Compared to the eastern Pacific, the vent faunas are less
varied; bivalve mussels appear to be uncommon, and
tubeworms absent (Gage and Tyler, 1991).
Mariana Trough
This back-arc spreading centre borders the subduction zone
of the Mariana Trench. It is isolated from the main mid-
ocean ridge system. The vent-fauna is very different from
those of the eastern Pacific, and is dominated by a sessile
barnacle (the most primitive living barnacle species known),
limpets and anemones. The giant bivalves of the eastern
Pacific are replaced by a large, hairy-shelled gastropod
(Gage and Tyler, 1991).
Shallow-water hydrothermal vents
Vents at depths of less than 20m have been described off
the Palos Verdes Peninsula, California. They support a
diverse assemblage of colourless chemosynthetic bacteria
similar to those of deep-sea vent sites, which form mats
around the vent openings. The mats provide nourishment
for the mollusc Haliotis cracherodii (Kleinschmidt and
Tschauder, 1985), commonly known as black abalone.
COLD SEEPS
Cold sulphide and methane-enriched groundwater seeps
occur near the base of the porous limestone of the Florida
Escarpment, as well as in the Gulf of Mexico (Fig. 10.4).
The seeps support a dense faunal community associated
with a covering or mat of bacteria on the sediment surface.
These communities are strikingly similar in taxonomic
composition to the hydrothermal vents of the east Pacific,
a fact which points to a common origin and evolutionary
history for both community types (Hecker, 1985). The
community consists of large mussels and the vestimentiferan
worm Escarpia laminata, as well as galatheid crabs,
serpulid worms, anemones, soft corals, brittle stars,
gastropods and shrimps. Mussel densities appear to be
linked to methane levels in the water, whilst tubeworm
density may be correlated with the hydrocarbon loading of
the sediment (Gage and Tyler, 1991).
Tectonic subduction zone seeps
Subduction seeps are more diffuse and lower in temperature
than hydrothermal vent seeps, and are rich in dissolved
methane. They are known to occur off Oregon, where the
fauna includes species of Lamellibrachia and large
vesicomyid bivalves, and in the Guaymas Basin in the Gulf
of California, where thick bacterial mats cover the sulphide
and hydrocarbon-coated sediment. The cold Japanese
subduction zone seeps occur at a depth of 1,000m in
Sagami Bay near Tokyo and in the subduction zones of the
trenches off the east coast of Japan. The communities vary,
but include dense benthic assemblages dominated by
Calyptogena clams associated with a stone crab Paralomis
sp., sepulid worms, sea anemones, galatheid crabs,
swimming holothurians and amphipods (Gage and Tyler,
1991).
Other colonised deep-sea seepage sites include a cold seep
to the east of Barbados dominated by the mussel
Bathymodiolus, vesicomyid bivalves and vestimentiferan
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101
1. Biological Diversity
worms. Dense communities, probably of recent origin, have
also been discovered on the Laurentian Fan on the south-
east Canadian continental margin. These include vesicomyid
and thyasirid bivalves, gastropods, pogonophorans,
galatheid crabs and bacterial mats. There is evidence that,
like hydrothermal vents, cold seeps are ephemeral and
cyclic. However, many species are now known to occur in
the deep sea in a variety of similar sulphur and other
compound-reducing habitats, and have been found to eccur
around such temporary habitats as oil-rich whale carcasses,
which may be important ‘stepping-stones’ for
chemosynthetic-dependent deep-sea animals (Gage and
Tyler, 1991).
References
Angel, M.V. 1982. Ocean trench conservation. Commission on
Ecology Papers No. 1. TUCN.
Angel, M.V. 1991. Biodiversity in the deep ocean. A working
document for ODA. Unpublished MS.
Buzas, M.A. and Culver, S.J. 1991. Species diversity and dispersal of
benthic foraminifera. Bioscience 41(7):483-489.
Gage, J.G. and Tyler, P.A. 1991. Deep-sea Biology: A natural history
of organisms at the deep-sea floor. Cambridge University Press.
Grassle, J.F. 1985. Hydrothermal vent animals: distribution and
biology. Science Vol.229.
102
Grassle, J.F. 1986. The ecology of deep-sea hydrothermal vent
communities. Advances in Marine Biology Vol. 23. Academic Press.
Grassle, J.F. 1989. Species diversity in deep-sea communities. TREE
4(1).
Grassle, J.F. 1991. Deep-sea benthic biodiversity. Bioscience 41(7).
Hecker, B. 1985. Fauna from a cold sulphur-seep in the Gulf of
Mexico: comparison with hydrothermal vent communities and
evolutionary implications. Biological Society of Washington Bulletin
6:465-473.
Hessler, R.R. and Sanders, H.L. 1967. Faunal diversity in the deep-
sea. Deep-Sea Research 14:65-78.
Kleinschmidt, M. and Tschauder, R. 1985. Shallow-water
hydrothermal vent systems off the Palos Verdes Peninsula, Los
Angeles County, California. Biological Society of Washington
Bulletin No.6.
Rex, M.A. 1983. Geographic patterns of species diversity in the deep-
sea benthos. In: Rowe, G.T. (Ed.), Deep-Sea Biology. Volume 8,
The Sea. John Wiley and Sons, New York. Pp.453-472.
Rowe, G.T. 1983. Biomass and production of the deep-sea
macrobenthos. In: Rowe, G.T. (Ed.), Deep-Sea Biology. Volume 8,
The Sea. John Wiley and Sons, New York. Pp.453-472.
Thorne-Miller, B. and Catena, J. 1991. The Living Ocean:
understanding and protecting marine biodiversity. The Oceanic
Society of Friends of the Earth, Washington DC.
Wolff, T. 1970. The concept of the hadal or ultra-abyssal fauna. Deep-
Sea Research 17:983-1003.
Wolff, T. 1977. Diversity and faunal composition of the deep-sea
benthos. Nature 267:780-785.
11. SOIL MACROFAUNA
Compared to conspicuously diverse habitats such as tropical
rain forest or coral reefs, the soil as a habitat in its own
right, with its own rich fauna and flora, is often
overlooked. It supports, however, a wide array of diverse
animals, with representatives from every major phylum in
the animal kingdom except the coelenterates and the
echinoderms (Wallwork, 1976).
SOIL AND SOIL FAUNA
The soil habitat is not a uniform environment. Examination
of a vertical section of the profile of a mature soil will often
teveal several layers reflecting its past history and
development. This sequence, from the organic litter layer
on the soil surface to the parent material below, can be
divided into four main horizons (Eisenbeis and Wichard,
1985):
O-horizon: organic upper layer of plant debris lying on
the surface of the mineral soil
A-horizon: upper, fine mineral soil permeated by
organic material
B-horizon: weathered, rough mineral soil coloured by
small deposits of humus
C-horizon: original, unweathered material.
The organic layer (O-horizon) can often be further
subdivided into three sub-layers: the leaf litter layer, the
fermentation layer and the humus layer, in a downward
succession (Wallwork, 1976). The actual depth that the O-
horizon attains is dependent on the rate of input from the
covering vegetation and the rate of decomposition.
The term ‘soil fauna’ can be used to encompass a large
number of animal species, including any which spend a
proportion of their life cycle in the soil, on the soil surface,
or in the leaf litter. The soil fauna contains numerous life
forms adapted to a great variety of microhabitats. In an
attempt to clarify the distribution of organisms within the
soil, Kevan (1962) proposed three categories, in terms of
their respective adaptations to life in the soil:
Euedaphon: inhabitants of the mineral soil, e.g. most
earthworms, all Symphyla, many mites
Hemiedaphon: inhabitants of the litter and fermentation
layer, such as many woodlice and millipedes
Epedaphon: inhabitants of the soil surface, such as most
ground-beetles and scorpions.
These categories are widely used, although some later
authors (e.g. Eisenbeis and Wichard, 1985) have modified
the definitions. Any given taxonomic group may include
species in more than one of the above categories, as well as
species which are not considered soil fauna.
Table 11.1 shows all taxonomic groups to be considered in
this context, with the term soil fauna being defined as
narrowly as practicable. Taxonomic level varies from
phylum to family. Taxonomic sequence follows Barnes
(1984).
103
Soil Macrofauna
Patterns of soil-fauna research
Although research is being conducted on the key soil
groups, much of it is limited to individual genera or species
rather than whole orders; and it covers only a few of the
major habitats. Thus, it is very difficult to build up a
picture of the total fauna of a region using literature
primarily on soil research.
The taxonomic precision of the primary literature also
varies through time. The 1940s and 1950s saw a peak of
species-level identifications by ecologists. Since the 1960s,
ecological and taxonomic interests have developed, so few
soil ecologists now provide species lists in their papers.
Data are now more often presented at order level, with
emphasis on biomass and productivity rather than on species
assemblages. Very recently, there have been moves to
revive taxonomic competence among ecologists
(Erzinclioglu, 1989; Dempster, 1991).
Other types of literature, such as general guides to animal
groups, identification keys, and taxonomic monographs,
provide useful information, but many are dated, thus
reducing the accuracy of their assessment of species totals
for a region. Many are also only the result of brief
collecting expeditions and so can only be considered
preliminary markers of the possible species richness.
As our knowledge of the soil fauna and habitat expands, so
our appreciation of its faunal diversity increases, sometimes
ten-fold. The estimated world total of Pseudoscorpiones
recently rose from 1,300 (Levi et al., 1968) to 3,000
(Davies et al., 1985), and of Collembola from 1,500-2,000
(Wallace and Mackeras, 1970) to 10,000-20,000
(Greenslade and Greenslade, 1983).
This is in line with the trend shown by invertebrate
diversity estimates in general. The degree to which current
figures for soil biodiversity may be relied upon is
geographically patchy: some areas have comprehensive and
up-to-date lists for most groups, and these have been
relatively stable for several decades despite an increasing
pace of ecological and biogeographic research (e.g.
Britain); a few others, such as Australia, are attempting to
produce comprehensive overviews; but in most countries,
the literature is becoming narrower and less easily used.
Ecological functioning and importance of soil fauna
The soil is basic to most terrestrial ecosystems, and the
health and functioning of the soil relies heavily on the
activities of soil fauna. The initial formation of soil, for
instance, at the end of a glaciation, depends greatly on
detritivores to help in cycling of nutrients and humus
formation. The accumulation of the latter is responsible for
the development of the soil through time. The role of soil
invertebrates in these pioneer phases must be considerable:
several groups of invertebrates are known, from the fossil
record, to have colonised newly exposed areas well in
advance of the vascular flora (Buckland and Coope, 1991).
The soil fauna is also a major vector of microorganism and
1. Biological Diversity
Table 11.1 Taxonomic distribution of soil macrofauna
PROPORTION OF GROUP PROPORTION OF EXTENT TO WHICH
WHICH ARE TERRESTRIAL SPP. SOIL SPP. UTILISE
TERRESTRIAL LIVING IN THE SOIL THE SOIL
Platyhelminthes:
Tricladida e ecco ecco
Nemertea e ecco eece
Nematoda eee eco ecco
Annelida:
Oligochaeta* eco eco eco
Mollusca:
Gastropoda eo ee e/ece
Crustacea:
lsopoda* ee
Amphipoda e ecc5e
Decapoda ® eo e
Chelicerata: Arachnida:
Scorpiones* ecco eoo0e eco
Pseudoscorpiones* eco5o eco eco
Uropygi ecoo ecc0e eco
Amblypygi ecoo ecco eco
Palpigradi ecco ecco ecco
Ricinulei e000 ecc5e ecco
Solifugae eooee ecco ee
Opiliones* eoce eco ee/eee
Araneae eee eo e
Acari*: i
Mesostigmata eooe eco eco
Prostigmata eo ee eco
Astigmata ecco e eco
Cryptostigmata ecoe eee rYy)
Onychophora ecco eco e
Uniramia:
Diplopoda* ecco ec0o ecco
Pauropoda eco0e ecco eoo5e
Chilopoda* ecc6e eco eee
Symphyla ecco ecoe ecco
Diplura eoc0e ecco eooe
Collembola* eco eco eco
Protura ecco ecco eooe
Thysanura coco ee eco
Embioptera* ecco eooe coe
Orthoptera*:
Gryllotalpidae eooe ecco eco
Tridactylidae ecco ecco ecco
Cylindrachetidae ecco eoo0o eco
Tetrigidae ecc5e ecco eco
Dermaptera* ecco eco ee/ece
lsoptera* ecco eco eoce
Blattaria* ecco eco ee
Psocoptera ecco eo °
Thysanoptera eoc0e eo ee
Homoptera eco ee eco
Coleoptera:
Carabidae * eco eco C ee/ece
Staphylinidae* eee eco ee/ece
Tenebrionidae eoo0e eco eco
Scarabaeoidea ecco coo eo
Elateroidea ecco eo eo
Cantharaoidea ecc5e ee eo
Hymenoptera:
Formicidae* ecco eo0o e/ee/ecee
Megaloptera e eo eo
Diptera eo ee ee
Notes: * indicates taxa considered key soil groups for which adequate biogeographic and taxonomic information has been located and which are
therefore considered in detail in this review. Nematodes are discussed in Chapter 9. Columns 2 and 3 are coded as follows: ©@@@ all species;
©©© most species; ©® some species; ® few species. Column 4 is coded as follows: @@ © obligate soil-dwellers; @@@ usually soil-dwelling, but
may at times climb vegetation etc.; @@ temporarily present, normally for a particular part of their life cycle (e.g. Diptera larvae); @ regular users
of the soil (i.e. for foraging) but able to spend much or all of their life in other microhabitats.
104
cryptogam propagules (Gerson and Seaward, 1977;
McCarthy and Healy, 1978). Many soil groups include
decomposers which are important in the breakdown and
recycling of nutrients throughout mature ecosystems. Most
temperate soils differ from soils at lower latitudes in having
a greater ‘standing crop’ of plant detritus, owing to the
lower decomposition rates, and tend to be deeper and with
a more elaborate profile, partly due to seasonality of
precipitation and the effects of frosts.
All of these processes, where soil invertebrates function as
pioneers and as facilitators of cycling in later seral stages,
are important in the rehabilitation of damaged ecosystems
where, for example, the vegetation and/or top-soil has been
lost. Earthworms have been shown to aid the development
of vegetation in derelict industrial sites; monitoring of their
population levels can therefore be used as an indicator of
the recovery of the habitat (Davis, 1986).
In most terrestrial habitats, the soil fauna is also important
for niche creation: that is, the activities of some groups
provide niches for other soil animals. Most importantly, the
actions of earthworms are largely responsible for the
structure of many soils, and their burrows allow access to
deeper parts of the soil not normally penetrable by other
groups; they thus provide retreats in the face of predation
or desiccation.
The soil macrofauna, in synergy with microorganisms, also
acts as a major link between the soil and non-soil habitats.
The role of these groups, and in catalysing the processes of
nutrient cycles, releasing minerals for uptake by vascular
plants, as well as providing a physical soil structure which
strongly influences the development of plant communities,
is crucial to the final appearance of the vegetation.
Many non-soil animals such as birds and mammals feed on
soil fauna regularly. For some groups, such as shrews
(Soricidae), hedgehogs (Erinaceidae), some wading birds
(Charadriiformes) and many reptiles and amphibians, soil
fauna may make up the bulk of their diet, at least during
part of the year.
PATTERNS OF SPECIES RICHNESS
This section contains a systematic account of the
biogeographic patterns of the key soil groups prefaced by
“notes on their biology. The higher level classification of
groups is not necessarily the same as followed elsewhere in
this volume. The reference list for data cited in the tables
below is available on request from WCMC.
Phylum Annelida
Sub-class Oligochaeta
The Oligochaeta is divided into two main soil-dwelling
groups, the earthworms (Lumbricina) and the potworms
(family Enchytraeidae). Oligochaetes are soft-bodied
segmented worms adapted to burrowing in the soil; they
include the only truly terrestrial annelids and are
ecologically a very important group.
Oligochaeta: Lumbricina
Earthworms are largely absent in highly acidic soils, such
105
Soil Macrofauna
as peatlands and heathlands: few species can tolerate a pH
lower than 4.0 (Wallwork, 1976). In base-rich soils,
however, they often constitute a high proportion of the total
animal biomass. Members of the family Lumbricidae
dominate the fauna in north temperate regions, and range in
size from less than lem to 35cm. Several other families
occur in warm temperate and tropical countries, the best
known being the Megascolecidae, some species of which
can exceed 3m.
Populations of 2.4-7.2 million earthworms per hectare have
been reported from rich permanent grassland habitats in
Britain (Cloudsley-Thompson and Sankey, 1968). They are
of considerable importance in soil processes (Sims and
Gerard, 1985). In addition to the benefits in agricultural
soils, earthworms are fundamental to the production of the
soil structure within which many other soil invertebrates can
live (Lavelle, 1983). Several important groups of soil fauna
are able to penetrate deep into the soil, and thereby survive
during dry weather, solely because of the network of fine
passages created by earthworms.
Oligochaeta: Enchytraeidae
The potworms comprise the terrestrial members of a family
many of whose members are freshwater or marine. They
are rarely more than 25mm in length and can tolerate acid
conditions much better than lumbricids; large populations,
of the order of thousands per m*, may be found in more
acid soils of oak woodlands and moorland peats (Wallwork,
1976). Few other groups are as successful in colonising the
rather sterile and water-logged soils of bogs; only
nematodes and Diplura (Eversham, unpublished) thrive
equally well in these conditions.
Areas which were covered by ice during the last glaciation
or which supported only tundra vegetation for several
millennia lost almost all their earthworms; post-glacial
recolonisation appears to have been restricted to a few
highly mobile, eurytopic species. Thus, Britain and north-
west Europe support a small fauna of only 10-30 species
and the earthworms of natural habitats are only slightly
more diverse than those of improved agricultural soils.
Countries bordering the Mediterranean have many more
species, with particular concentrations in the Iberian
Peninsula and Italy; even the French fauna reaches 97
species (Lavelle, 1983). In such areas, agricultural
improvement displaces the more stenotopic species. A
similar pattern is found in North America. Of nearly 400
lumbricid species recognised, only 5% occur in the northern
areas which were overlain by ice sheets. A whole
superfamily, the Crilodriloidea, is now confined to a small
area of the southern USA (Sims and Gerard, 1985).
There is limited evidence that the centres of species richness
in the southern hemisphere are now being threatened by the
introduction of north temperate species, which are
associated with agricultural soils but may be able to out-
compete the indigenous fauna (Ljungstrom, 1972).
Other aspects of family level distribution throw light on
much more ancient geomorphological history. A
consequence of the two effects (relict Gondwana
distributions and Pleistocene glacial defaunation) is seen,
for instance, in the much richer earthworm faunas of
1. Biological Diversity
southern hemisphere islands like New Zealand (192 species;
Lee, 1959) compared with that of a similar-sized landmass
such as Britain (28 species), which should for reasons of
island biogeographic theory be expected to acquire species
much more readily from its nearby continent. The
impoverished fauna of Iceland (8 species; Lavelle, 1983) is
an even more extreme example of this; it is likely that the
whole of this fauna is recently introduced by man. The
position of Japan in relation to the Eurasian landmass is
reflected in its relatively rich fauna (75 species; Easton,
1981).
Table 11.2 Soil species: Oligochaeta
TOTAL SPECIES 1,200
Lumbricina
New Zealand 192
France 97
Japan 75
Oregon (USA) 267
UK 25
Little Carpathians (East Europe) 22
Denmark 19
Sweden 13
Washington (USA) 137
Iceland 8
Enchytraeidae
North America 143
Europe 111
Little Carpathians (East Europe) 24
Phylum Crustacea
Almost all the terrestrial Crustacea belong to the order
Isopoda, the familiar woodlice, slaters or sowbugs. A very
few members of the mainly aquatic Amphipoda and land
crabs have also adopted a terrestrial existence.
Class Malacostraca: Isopoda
Some genera rarely venture up to the soil surface, whereas
others spend most of their existence among leaf litter and
grass roots, and a few forage regularly among herbaceous
vegetation or even in the lower branches of trees (Sutton,
1980). In all these cases, the major part of woodlouse diet
is probably dead plant matter, though some species have
been observed browsing on the living foliage of trees.
Isopods have developed a wide range of behavioural
adaptations to avoid desiccation.
Eurasian Isopoda appear to have a strong centre of diversity
around the Mediterranean - especially in Spain, Italy and
North Africa; it has been suggested that the fauna in these
areas is even more diverse than that of tropical sub-Saharan
Africa (P.T. Harding, pers. comm.). A few north-west
European species have been widely spread by man, and
make up a large proportion of the common synanthropic
woodlice of North America and other temperate regions,
and a few species are now almost cosmopolitan, e.g.
Ojlisticus convexus (Harding and Sutton, 1985). Although
superficially amongst the better-known soil macrofauna,
some woodlice are very small and cryptic, and occupy
narrow and obscure niches: for example, the coastal
shingle-bank fauna of northern Europe is only just being
106
discovered. In well-worked regions, there is clear evidence
of this niche specialisation, as well as landscape-scale
differentiation (Harding et al., 1991).
Some species associated with ancient natural habitats are
now threatened by agricultural change and other human
modifications of the landscape, such as the clearance or
replanting of ancient woodland, the drainage of wetlands
(e.g. Ligidium hypnorum in England), and general
disturbance of coastal habitats (e.g. Armadillidium album
throughout its range).
Table 11.3 Soil species: lsopoda
France 156
North America 100
Germany 60
UK 42
Holland 35
Little Carpathians (East Europe) 28
Phylum Chelicerata
Class Arachnida
Second only to the insects among the arthropods in terms of
species numbers, the arachnids are an ancient and mainly
terrestrial group.
Arachnida: Scorpiones
A morphologically rather uniform group of nocturnal
predators with modern species ranging from 1.3cm to 18cm
in length, but some Carboniferous species attained a length
of 86cm (Barnes, 1980). Although usually thought of as
typical of arid regions, there are many species which
require a humid environment and occur in tropical moist
forests. Most species occur in warm regions, but a few
occur near the snow-line in mountains, and a single species
occurs as far north as Canada (Levi et al., 1968).
Table 11.4 Soil species: Scorpiones
TOTAL SPECIES 3 1,000 +
Africa (S) 159
Iran 36
USA 20-30
Kenya (N) 26
Arabia 23
Tanzania/Kenya 23
Africa (W) 17
lraq 15
Israel f 15
Australia (W) 11
Syria 11
Turkey 11
Egypt 9
Israel (N) 8
Trinidad 7
Libya 6
Madagascar 6
Kenya (S) 6
Jordan 5
Tobago 3
It is possible that the ranges of some temperate species are
currently not entirely climatically determined, but reflect
incomplete recolonisation after the glaciation. From the
literature it appears that scorpions have a surprisingly even
distribution of species richness in the warmer regions of the
world. There is no evidence of any areas of marked
radiation, which may be characteristic of an ancient and
morphologically conservative group. The only exception is
the apparent radiation of the rich southern African fauna,
which parallels the high diversity of certain plant groups,
especially Erica and Protea (Good, 1964). An additional
factor in producing the even distribution of species, with
few areas of very high diversity, may be their mode of life:
they are bare-ground active hunters of large invertebrates
and small vertebrates, and consequently occupy a broad
niche space which cannot easily be partitioned between
species, even if individual population density is high. In this
respect, they provide an interesting contrast with the
Carabidae, another group of surface-active generalist
predators as discussed below.
Arachnida: Pseudoscorpiones
These small arachnids, the largest being only 8mm long and
most only 2-3mm, superficially resemble scorpions. Most
species live in leaf litter, and require a high humidity; in
suitable woodlands, very high densities may be attained,
with over 500 per m? commonly recorded, and peaks of
over 900 per m? reported (Gabbutt, 1967). The efficient
dispersal of species, particularly those with narrow
microsite requirements (nests, barns, decomposing
vegetation), is enhanced by phoresy - attaching themselves
to other arthropods, especially Diptera and occasionally
Coleoptera, and remaining attached until the host reaches
another patch of suitable habitat. The maximum diversity of
pseudoscorpions is widely believed to be in the tropics
(Wallwork, 1976), but the available data are very patchy,
with no comprehensive regional reviews, even in generally
well documented areas such as Europe and North America;
almost all the literature focuses on individual genera or
species. The classification at species and family level is still
in a state of flux. There is some evidence of microsite
specialisation at the landscape scale (Legg and Jones, 1988),
but few sites support a particularly rich range of
pseudoscorpions.
Table 11.5 Soil species:
Pseudoscorpiones
TOTAL SPECIES 3,000
North America 200
South Africa 109
Australia 99
Brazil 40
UK 26
Arachnida: Opiliones
The harvestmen or harvest-spiders have an average body
length of 5-10mm, but the largest tropical species reach
20mm with a leg length of 160mm (Barnes, 1980).
Harvestmen are abundant in leaf-litter and low vegetation in
most habitats. In some tropical forests, the predatory force
of harvestmen is thought to exceed that of the spiders
(Dalingwater, 1983), although they will also scavenge on
107
Soil Macrofauna
dead animals, and will eat a very wide range of organic
matter, including fruit. Many species forage on the trunks
and branches of trees when adult, but even these species
tend to spend most of their juvenile life in litter or grass
roots. Many species are nocturnal, as an adaptation to
avoiding desiccation; the more resistant species are able to
be active by day (Todd, 1949).
There are two ecological/systematic divisions in the
harvestmen which show contradictory distribution trends,
although the overall pattern is of higher diversity in the
tropics. The actively predatory Laniatores are almost
exclusively tropical and can be regionally diverse, e.g. 581
species in South America (Lawrence, 1931). The other
main group, the scavenging Palpatores, show the reverse
trend, with high diversity in the temperate regions: the
known South American fauna contains a mere 29
Palpatores, while Europe possesses 215 species, compared
to the meagre 14 native south-European Laniatores, of
which only eight occur in central/northern Europe
(Lawrence, 1931; Martens, 1978).
Harvestmen are more sensitive to desiccation than most
arachnids, so are ill-adapted to a desert environment. This
may explain the low diversity in Australia, for instance
(where there is a rich spider fauna), compared with the rich
Opiliones fauna in moist tropical forests, where they may
be able to out-compete the spiders (Dalingwater, 1983).
Table 11.6 Soil species: Opiliones
TOTAL SPECIES 3,500
South America 581
Europe 232
Africa (excluding S Africa) 201
New Zealand 170
Europe (N of Mediterranean) 110
North America 104
South Africa 90
Madagascar 69
China 60
Australia 38
UK 23
Holland 21
Arachnida: Acari (Oribatei)
There are seven major groups of mites and ticks in this
huge order, but only mites of three suborders occur
predominantly in soil. They have a worldwide distribution.
The Cryptostigmata or Oribatei are generally saprophagous,
some feeding directly on decomposing litter fragments while
others eat the fungi and bacteria which coat the litter. The
Mesostigmata also include some saprophages, but many
species are predatory. Prostigmatid mites are very varied in
form and habit and include many non-soil-dwelling species.
Of these three suborders the Cryptostigmata or Oribatei are
by far the best known, in terms both of taxonomy and
ecology; the prostigmatid and mesostigmatid soil-mites have
received less attention and are less confined to the soil.
Most Oribatei are less than 1mm long, some very much
smaller. High population densities can occur, with figures
of 130,000 per m? being unexceptional. They occur in a
wide range of soil and litter microhabitats the world over
1. Biological Diversity
(Luxton, in prep.) and play an important part in litter
decomposition, both directly (those which feed directly on
litter will consume about 20% of their body weight in litter
each day), and indirectly (they stimulate microbial action in
the litter). As the single most important fungivorous group
in the soil, up to 50% of microfungal grazing and spore
dispersal is attributable to oribatids (Eisenbeis and Wichard,
1985).
The available numerical data suggest that temperate soils
support a more diverse cryptostigmatic mite fauna than the
tropics, but this is almost certainly an artefact of sampling.
The British Isles, with 300 species, would appear to have
the richest concentration of any area, but also has the only
up-to-date checklist (Luxton, in prep.). The available world
literature concentrates almost entirely on generic and
species taxonomy, or the fauna of very small sampling
areas within atypical habitats. Oribatids are such an
important group within the soil, being geographically and
biotopically ubiquitous, that their overall biodiversity
pattern will be of great interest when sufficient comparable
data have accumulated.
Table 11.7 Soil species: Acari
(Oribatei)
TOTAL SPECIES 7,000
UK 300
USSR (European) 278
Bulgaria 250
Japan 170
Arctic 144
Little Carpathians (East Europe) 129
Canada (N) 106
Peru 91
Ghana 52
Alaska 10
India (8 )
Phylum Uniramia
Class Chilopoda
This class comprises the centipedes, an important group of
elongate, swift and agile predators which play a
considerable part in most soil ecosystems. The class may be
divided into four orders, representing the four main lines of
morphological adaptation: the Geophilomorpha, the most
subterranean group of centipedes, rarely seen on the
surface; Lithobiomorpha;Scolopendromorpha, including the
largest of all centipedes, some reaching almost 30cm in
length; Scutigeromorpha, the majority of which live in dry,
rocky habitats, hunting among rocks and scree. Several
families of centipede are better represented at lower
latitudes, the Scutigeromorpha in particular being confined
to warm-temperate and tropical regions, though a few
species occur inside human habitations further north. The
trend in family distribution appears, from the very limited
data, to be reflected in species richness too; but the
accessible literature on centipedes is fragmentary, and even
the most thoroughly researched areas such as northern
Europe still have many areas of taxonomic confusion.
108
Table 11.8 Soil species: Chilopoda
TOTAL SPECIES 3,000
Peru 74
Germany 60
Transvaal 47
Natal-Zululand 42
UK 41
Holland 35
Canada (N) 29-31
Africa (SW) 27
Congo 10
Bermuda 7
Tunisia 7
Cyprus 6
Peru (NE) (6 )
Arctic 3
Class Diplopoda
The millipedes live in litter, under bark, and in the soil,
being active in the open only after dark. Some are cave-
dwelling, and several species live commensally in the nests
of ants. All millipedes are predominantly saprophages,
feeding on dead leaves, fallen logs and branches of trees,
though some may also occasionally browse on mosses,
lichens, algae or even living vascular plants. They often
occur at high densities, and can be the main shredders of
leaf litter in woodland soils that are too acid to support a
rich earthworm population (Blower, 1985).
Documented diversity is rather low in most areas, including
tropical Africa, but there is a high figure for North
America. This suggests a Nearctic warm-temperate peak of
diversity, enhanced by the absence of east-west
geographical barriers in the Americas (where the main
mountain ranges run north-south). This may have permitted
much greater northward spread of taxa than in Eurasia
(where there are major physical barriers - Pyrenees, Alps,
Himalayas etc. - running east-west, and restricted post-
glacial recolonisation of the region).
Table 11.9 _ Soil species: Diplopoda
TOTAL SPECIES 7,000
Central America and Mexico 750+
North America 749
France 250
Natal-Zululand 188
Germany 160
Peru (NE) 78
Transvaal 69
Congo 67
Madeira 53
UK (1958) 52
Holland 45
Denmark 39
Little Carpathians (East Europe) 31
Africa (SW) 17
Bermuda 8
Cyprus 6
Tunisia (1)
The four-fold difference in recorded diversity between
Britain (41 species) and Natal-Zululand (188 species) may
be partly owing to the glacial effect; but millipedes are
considered to be largely woodland/forest animals. Southern
Africa has supported much more extensive woodlands with
a stable history, throughout the Quaternary. The very low
diversity in the Arctic probably reflects the low primary
productivity, and thus the limited vegetable detritus for
millipedes to consume. An extreme example of island
speciation in soil fauna because of natural barriers may be
found on Madeira, where 25 of the 53 species are now
considered endemic. Ecological segregation in those regions
which have been adequately studied tends to be on a
macrohabitat scale, with grassland, woodland or sand-dune
species, for instance, rather than intensive multispecies
resource partitioning within a single habitat (Blower, 1985).
Class Oligoentomata: Collembola
The collembolans or springtails are small apterygotes
(primitive insect-like hexapods; Dohle, 1988), seldom
greater than 5mm long.
Like the mites, they have a cosmopolitan distribution,
Tanging from the seashore to high mountain-tops, and from
the equator to the poles. Similarly, they can occur in very
high densities, the smaller species reaching hundreds per
cm? in ideal conditions. Species vary in their desiccation
tolerance, so that different microsites in a habitat will
support different species. The majority of Collembola are
saprophages, feeding on decomposing plant and animal
debris, although a few are predators, and others are small-
scale pests of crops, notably the Lucerne Flea Sminthurus
viridis. Because of their enormous densities and ubiquity,
springtails are a crucial food-source for many small soil
predators, including pseudoscorpions, and some staphylinid
and carabid beetles.
Like the Oribatei, the Collembola appear to show a trend to
higher diversity in temperate regions than in the tropics.
Again, data quality may be suspect, but appears to be
considerably higher and more uniform for Collembola than
for Oribatei. A possible explanation of this trend proposed
by Rapoport (1982) is that temperate soils are richer in
nutrients and organic matter, as well as being more
elaborately structured.
Comparing similarly-sized land masses with broadly similar
climate reveals a constancy of collembolan fauna: Britain
(300 species), Japan (241 species) and New Zealand (293
species) (Chinery, 1973; Rapoport, 1982). However, it is
almost certain that the majority of species have yet to be
found: for instance, Wallace and Mackeras (1970) could
Tefer to only 215 described species, whereas Greenslade and
Greenslade (1983) estimated there were 1,000-2,000
Australian species. The figures for mainland North
America, lower than for Britain, are likely to be a sampling
artefact.
Little has been published on patterns of collembolan
endemism, but many species and genera have wide
geographic ranges, implying some effective mechanism for
long-distance dispersal, possibly wind-blown or rain-blown
eggs. The available figures for the Tasmanian fauna
contrast the native forest fauna, where up to 40% of species
109
Soil Macrofauna
are endemics, with that of managed grassland, where only
1-2% of species are endemic and the majority are
cosmopolitan (Greenslade and New, 1991). This clearly
suggests that the Collembola will be highly sensitive to
human impacts on natural and semi-natural vegetation;
unfortunately, little research has been done elsewhere in the
world.
Table 11.10 Soil species: Collembola
TOTAL SPECIES 10,000-20,000
Australia 1,000-2,000
UK 300
USSR 300
California (USA) 150
Little Carpathians 143
(East Europe)
Peru 97
Arctic 91
Iceland 58
Philippines 37
Sudan 24
Class Pterygota: Dermaptera
The earwigs are a distinctive order of medium-sized insects
allied to the Orthoptera. Although often hiding among litter
or in the soil during the day, many species forage
nocturnally among vegetation, flying readily and climbing
trees (Imms, 1957). They are included here as soil fauna
because almost all return to the soil to breed. Most species
are thought to be omnivorous (Marshall and Haes, 1988).
Earwigs are essentially tropical and subtropical in
distribution. Most species are sedentary, so individual
species tend to have rather small geographic ranges, and
consequently the fauna of each region contains a high
proportion of endemics. The African fauna has been more
intensively studied than others. Central Africa appears to be
an important centre of diversity, particularly for the more
primitive families of earwigs; it contains about 30% of the
known world species of the ancient Carcinophoridae, for
example, but only 18% of the more advanced Labiidae
(Brindle, 1973). Literature on other tropical regions is
sparse, although the Indian subcontinent appears, like
Africa, to hold important concentrations of species.
Table 11.11 Soil species: Dermaptera
TOTAL SPECIES 1,200
Africa 298
India 185
Australia 60
USSR 26
USSR (European) 17
California (USA) 10
UK 5
Iceland 1
Only a very few earwig species are truly cosmopolitan,
although their lifestyle makes them susceptible to accidental
transport through commerce. Many such casual
translocations lead only to temporary establishment, but if
the climate is suitable a species may become more
1. Biological Diversity
widespread. For example, the Indo-Australian species
Marava arachidis is now well established in Africa and the
Americas, but occurs only sporadically in Britain and
northern Europe, usually in warehouses of imported organic
materials (Brindle, 1973; Marshall and Haes, 1988). The
sole cosmopolitan temperate species, the European Forficula
auricularia is the common garden earwig in North America
and elsewhere, though in the tropics it occurs mainly in
montane areas. It has been implicated in the demise of three
endemic earwigs of the genus Anisolabis in Hawaii
(Howarth and Ramsey, 1991).
Pterygota: Embioptera
This primitive order comprises small to medium-sized soft-
bodied cylindrical insects, commonly known as web-
spinners. The females of most species are believed to be
predominantly herbivores, while the males’ diet may
include other insects and soil arthropods.
Web-spinners are essentially tropical animals. The small
numbers of European species are confined to the south,
their northern limits being the Crimea, Bulgaria and the
shores of the Mediterranean, although a few species occur
further inland in Spain. The American fauna totals over 70
species, of which three are introduced and the rest are
endemic (Ross, 1944). The highest concentration of species
is probably in Australia (65 species (Ross, 1970)).
Overall, web-spinner species occur in widely-scattered,
isolated areas, the group distribution being highly
discontinuous.
Table 11.12 Soil species: Embioptera
TOTAL SPECIES 100
Australia 65
South America 44
Europe and Mediterranean 24
Central America 15
USA 12
Europe (S) 5
USSR 2
California (USA) 3
USSR (European) 1
Pterygota: Orthoptera
Four families of Orthoptera are largely soil-dwelling: the
Gryllotalpidae, Tridactylidae, Cylindrachetidae and
Tetrigidae. Many other species of grasshoppers and crickets
spend some of their time among leaf litter and/or lay their
eggs in the soil but are not included here because a
significant part of their life-cycle takes place away from the
soil.
The literature on the three soil-dwelling groups of
orthopteroids is partial and fragmented. It is thus difficult
to draw global conclusions at this stage.
Orthoptera: Gryllotalpidae
The mole-crickets are a small and specialised family of
large, bulky insects which construct burrows mainly for
feeding. Although found mostly in natural grasslands, they
occasionally reach pest status by attacking root crops,
110
especially (in temperate areas) potatoes (E.C.M. Haes,
pers. comm.). Most species can fly, and can therefore
colonise new areas. They are rare in cool-temperate regions
and more diverse in warm-temperate ones.
Many species are phenotypically very similar, but there
may be genetically-isolated cryptospecies awaiting
recognition. Those species already described are fairly
uniformly distributed between the main biogeographic
regions, with no marked concentrations apparent from the
literature. A few species are occasionally transported by
man, mainly among root-crops; and the commonest
Eurasian species, Gryllotalpa gryllotalpa has been
introduced into North America.
Table 11.13 Soil species: Gryllotalpidae
TOTAL SPECIES 50
Australia 7
USSR 3
USSR (European) 2
UK 1
Orthoptera: Tridactylidae and Cylindrachetidae
The pigmy mole-crickets are not closely related to
Gryllotalpidae, but have converged on the same lifestyle
and acquired the same modifications of body form. They
are relatively small - less than 10mm long - and live in
damp sandy soils usually close to water.
The Tridactylidae are widely scattered in warm-temperate
and subtropical regions, whereas the Cylindrachetidae are
confined to Australia, New Guinea and Patagonia (Imms,
1957); the latter probably indicative of the family’s early
evolutionary origins on Gondwanaland.
Table 11.14 Soil species: Tridactylidae
and Cylindrachetidae
TOTAL SPECIES 50
Australia 4
USSR 4
USSR (European) 3
Orthoptera: Tetrigidae
The groundhoppers or grouse-locusts are relatively small,
usually less than 20mm. Most are found in damp
microsites, such as river or pond margins. The eggs are
often drought-resistant, enabling species to occupy
seasonally-wet habitats (Hartley, 1962). Most species are
unable to fly.
The Tetrigidae is a large group, with many described
species. They appear, from the limited figures available, to
be best represented in warmer regions, the Australian fauna
being among the largest, though quite high concentrations
have been described in some cool temperate areas.
However, their taxonomy is still being clarified, and the
ecological distinctions between closely-related species are
only just beginning to be determined, even in western
Europe (Devriese, 1990).
Table 11.15 Soil species: Tetrigidae
TOTAL SPECIES 700
Australia 70
Europe, Asia and N Africa 50
USSR 14
USSR (European) 9
UK 3
Class Pterygota: Blattaria
Small (e.g. temperate Ectobius, 5-7mm) to large (e.g.
tropical Blaberidae, up to 15cm) insects. Most of the world
fauna lives in low vegetation or on the ground, probably as
scavengers; dense populations can occur in the litter layer
of warm forests, and some species occur in caves. A
handful of cosmopolitan species are pests and can be very
abundant in domestic situations.
Cockroaches are characteristic of tropical moist forests,
which support the largest diversity. Australia, for instance,
has 439 species, most found in the native forests.
Comparing two areas of roughly equal size, the British Isles
support only three species, all in the genus Ectobius,
whereas the West Indies are home to 156 species, including
representatives of all the major families. There are several
cosmopolitan species spread by man and now established in
most countries. For this reason, published checklists,
particularly in colder regions, often overestimate the
indigenous fauna by including aliens which are restricted to
heated domestic premises, e.g. Britain has three native and
23 casual or introduced species, of which five are well
established (Marshall and Haes, 1988); the whole of the
USSR has 41 native and 12 alien species.
Table 11.16 Soil species: Blattaria
TOTAL SPECIES 3,500
Australia 439
Africa (W) 300
West Indies 156
USA 55
USSR 50+
California (USA) 5-6
UK 3
Pterygota: Isoptera
The termites or ‘white ants’ are one of two main groups of
soil-dwelling social insects (the others being the ants,
Hymenoptera: Formicidae) whose colonies consist of a
complex caste system, in which four main types can be
recognised: the queen(s), workers, soldiers, and alate
sexuals. The most primitive types are wood-boring and
feeding, making no external modification to the decaying
timber in which they live; such forms generally lack the
worker caste. Certain genera may become pests by boring
in domestic timbers. The remaining families are more
exclusively soil-dwelling, some simply excavating galleries
underground with little surface protrusion, while others
construct large termite-mounds or termitaria which extend
111
Soil Macrofauna
the nest many metres above the soil surface and form a
conspicuous feature of the landscape in African and
Australian scrub-grasslands. Many species feed in the same
manner as earthworms, ingesting the soil detritus,
microfungi and bacteria, or upon the roots of grasses and
other plants. Others cultivate elaborate ‘fungus gardens’ on
compost pre-prepared from vegetable matter. The majority
of these more advanced species do not forage beyond the
confines of the nest, unlike social Hymenoptera. The actual
impact of termites on tropical ecosystems is still being
evaluated (e.g. Collins, 1980, 1983, 1989).
In addition to their direct contribution to biodiversity,
termites are important in providing niches for an extensive
cohabiting fauna in their nests, ranging from commensals to
symbionts, parasites and specialist predators.
Termites occur widely outside the polar and cold-temperate
regions, except in the Palaearctic. The Ethiopian region
appears to possess the richest diversity of genera as well as
species, and is thus probably the most important centre of
termite evolution (Bouillon, 1970). It contains the largest
proportions of endemics. High numbers of species are also
found in South America and the oriental region.
Broad patterns of temperature explain much of the variation
in termite diversity. In the northern hemisphere, a strong
correlation between diversity and latitude has been found
(Sutton and Collins, 1991), though this may be a slight
over-simplification: the correlation would be far less clear
using southern-hemisphere data, because of the rich termite
fauna of Australia, which extends beyond the Tropic of
Capricorn.
Table 11.17 Soil species: Ilsoptera
TOTAL SPECIES 2,000
Ethiopian Region 570
South America 499
Oriental Region 434
Australia 182
Congo and Cameroon 78
Thailand 74
Palaearctic Region 41
Myanmar 39
Pakistan (W) 30
California (USA) 15
Mexico(W) 15
New Zealand 11
USSR 4+
Europe 2
Pterygota: Hymenoptera (Formicidae)
The ants are morphologically conservative but behaviourally
diverse social insects with an elaborate caste system. Their
nests vary from a few individuals in a space of less than
lcm? contained insidt a dead twig (e.g. Leptothorax) to
huge soil-based mounds with hundreds of thousands of
foraging workers, which may be the dominant predatory
force in whole forests (Brian, 1977). The diversity of
individual size and feeding ecology allows many species to
coexist in an area, and to partition resources, thereby
avoiding competition (Davidson, 1978). The majority of ant
nests are situated either within the mineral soil, or in the
1. Biological Diversity
litter layer; although with deserved reputations as predators,
many species also consume large volumes of plant material,
especially seeds. Quite a high proportion of ant species have
complex interactions with other ants. Like termites, ants
also interact elaborately with other invertebrates, thereby
increasing invertebrate diversity, through providing a range
of additional niches within their nests; the range of
symbiotic, inquiline, commensal, scavenging, parasitic and
predatory lifestyles closely parallels those found within
termite nests.
The ants are a large and diverse group, with most species
in tropical regions, and a sharp decline toward the cool-
temperate. Even on a small scale, in Europe and North
America, there is a clearly marked latitudinal decline in
diversity (Cushman and Lawton, in press); for instance,
France has 180 species, whereas Britain has only 46
including introductions. The Palaearctic and Nearctic faunas
are roughly equal in total diversity and pattern of species
richness, their post-glacial colonisation apparently being
unaffected by the topographic differences between the
continents described under Diplopoda. This may be because
the winged queens of ants are highly mobile, and so could
travel long distances and recolonise virgin habitats as they
became available with the retreat of the ice-sheet. This
could also be the reason why Britain (46 species) has twice
as many species as New Zealand (23) despite the fact that
the total fauna of Oceania is much richer than that of
Europe: the isolation of New Zealand is too great for
uncontrolled flight to convey large numbers of species.
Ants have been the focus of much ecological research and
speculation over the past 40 years. It has recently been
observed that in Europe and temperate North America there
is a latitudinal cline in individual mean size, with larger ant
species in the boreal forest and many more tiny species
arcund the Mediterranean/southern USA (Cushman and
Lawton, in press). Further explanations of regional
biodiversity have been related to vegetation patterns
(Greenslade and New, 1991), and Australian work has also
shown a high species turnover (beta diversity) across the
continent.
Table 11.18 Soil species: Hymenoptera
(Formicidae)
TOTAL SPECIES 10,000
Neotropical Region 2,233
Australia 1,100
North America (+USA) 585
USA 400+
California (USA) 200+
France 180
Sweden 61
Denmark 49
Finland 47
Norway e 46
UK 46
New Zealand 23
Pterygota: Coleoptera (Carabidae)
The ground-beetles and tiger-beetles may be the largest of
all families in terms of total species; over 40,000 species
are described (Erwin et al., 1979). They are ecologically
very wide-ranging, in diet varying from obligate herbivore
and detritivore to highly specialised predator. Their size
ranges from less than 2mm to several centimetres, and they
occupy almost all habitats from permanently waterlogged
soils to the driest deserts. Although a proportion of forest
species forage in the canopy, and rest under bark, the great
majority are closely linked to soil and litter. Ground-beetles
can reach high diversity in small habitat patches because of
the variety of ways in which they can divide up the food
resource, microsites, and time (different species being
diurnal, nocturnal or crepuscular) (Greenslade, 1963).
With over 40,000 described species, the Carabidae are
potentially valuable in analysing patterns of soil fauna
distribution. Unfortunately, many areas still lack
comprehensive reviews of their fauna, so the available
literature remains patchy. However, the high diversity
reported from the main tropical landmasses is probably a
genuine effect; these areas did not suffer the extremes of
recent glaciations, and the long periods of stability may
have allowed local speciation to occur.
One of the most striking examples of intensive local
speciation is provided by the tiger-beetles (sub-family
Cicindelinae) in India, where there are 150 species in the
genus Cicindela. The explanation of this high diversity is
probably a complex of past dispersal, ecological isolation
(largely through local climatic effects) and habitat
specialisation (Pearson and Ghorpade, 1989). This contrasts
with the low diversity of other surface-dwelling generalist
predators such as scorpions.
There is a rich boreo-montane fauna in the northern
hemisphere: carabids make up a large proportion of most
European early post-glacial fossil deposits (Atkinson, Briffa
and Coope, 1986), and this highly mobile element is equally
important in North America - hence the rich
Canadian/Alaskan fauna (850 species, Lindroth, 1969). The
comparison of Britain (350 species) with New Zealand (538
species, Hudson, 1934) probably reflects local speciation on
the oceanic island: over 90% of New Zealand’s terrestrial
arthropods are endemic (Howarth and Ramsey, 1991). In
contrast, Britain has in effect only been partially recolonised
from mainland Europe because of the breach of the land
bridge to Europe by the English Channel, and has only a
single ‘endemic’ carabid, Tachys edmondsi (Lindroth,
1974).
Table 11.19 Soil species: Coleoptera
(Carabidae)
TOTAL SPECIES 40,000
Neotropical Region 5,000
North America 2,500
Australia 1,613
California (USA) 800
New Zealand 538
UK 350
Iraq 176
Although tropical forests support a very rich carabid fauna,
arid grasslands are less rich; this is in part because of their
replacement by the more drought-adapted Tenebrionidae.
For example, whereas Britain has a mere 44 tenebrionids in
a beetle fauna of over 3,000 species, Morocco has 711
species, which amounts to 15% of the total fauna (Kocher,
1958).
Pterygota: Coleoptera (Staphylinidae)
The rove-beetles range in size from less than 1mm to
several centimetres. Many species are predatory, but others
feed on decaying organic matter - vegetation, dung or
animal corpses. A number of species occur in ants nests,
some commensally or scavenging, others partially predatory
on the ant brood, but often providing the ants with a sweet
secretion in return. As a group, the Staphylinidae are an
important predatory force in moist temperate habitats
(Hammond, in prep.), perhaps rather less so in the tropics.
Many species are difficult to identify, and they are therefore
often excluded from surveys.
The rove-beetles are less well-known than the carabids, but
the existing numerical data reveal several patterns among
the temperate fauna. Most noticeably, the staphylinids
outnumber the carabids in each documented area in the
northern hemisphere, whereas in the southern, the reverse
is true. One possible explanation for this is that rove-beetles
are more prone to flying and were thus able to continue
colonising new areas despite rising sea-level after the last
Ice Age. The lower diversity in the southern hemisphere is
harder to explain, and data are too few to evaluate with
confidence; in some cases (e.g. Australia, with only 650
species) the generally more arid climate may limit the
Staphylinidae.
Table 11.20 Soil species: Coleoptera
(Staphylinidae)
TOTAL SPECIES 27,000
North America 2,800
California (USA) 1,000
UK 1,000
Australia 650
West Indies 468
Morocco 423
New Zealand 216
GENERAL PATTERNS OF DIVERSITY
This preliminary study has shown that the different groups
of soil macrofauna function ecologically in very different
ways and that most trends in distribution will be group-
specific. The soil fauna is such a diverse group that the
distributional trends within, for example, scorpions may run
counter to those of the Collembola. In a more detailed
study, it may thus be better to consider the major groups
separately: the differences between soil groups may be
greater than those between soil and non-soil members of the
same group.
It would thus be an over-simplification to look for a single
pattern of soil faunal biodiversity. That said, there are some
indications of global pattern which hint at concentrations of
species very different from those found in most plant and
animal groups.
113
Soil Macrofauna
The usual trend towards higher diversity in the tropics
compared with temperate regions is certainly apparent in
some soil groups such as the scorpions, solifugids and
Orthoptera. However, the limited information available for
others, such as the Collembola, appears to show the
reverse: temperate faunas may be more diverse than tropical
ones. A possible explanation lies in the difference between
the profiles of the two soils; tropical soils do not possess
the depth or varied horizons seen in temperate ones. This
is because of efficient re-cycling processes producing a low
organic content, and lack of thermal seasonality (Rapoport,
1982). Both of these factors reduce the niche space and
habitat quality of the soil, and consequently the soil-fauna
diversity that it can support. A more fundamental difference
is revealed when the respective ages of the soils are
considered. The older tropical soils, such as those in
Australia and Africa, are strongly leached and weathered,
while the temperate soils, such as those in northern Europe,
possess large areas of unweathered rock left by the
retreating ice-caps of the last glaciation. The latter therefore
have a higher mineral content, and a steady release of
inorganic nutrients, which enhances the fertility of the soil.
It is premature to identify centres of diversity and
endemism with any confidence although a few areas on
present evidence stand out. The faunas of South Africa,
Australia, and the Mediterranean Basin are richer than the
average in most groups. That of New Zealand shows a
higher degree of endemism than other similar-sized areas,
and is species-rich in some groups such as the Carabidae.
In many, the South American fauna is too poorly described
in the literature to allow detailed comparison, but the few
available figures suggest it is very rich in many groups.
Explanations of patterns of diversity depend on several
different effects, which may be contradictory. For example,
post-glacial history may have led to an impoverished fauna
in large parts of the northern hemisphere, yet it is also
responsible for the elaborate soil structure and landscape
mosaic seen in many areas of Europe and North America,
which enhance diversity. These two effects are jointly
responsible for the Mediterranean species concentrations in
several groups: during the glaciation, large numbers of
species appear to have survived in Mediterranean refugia,
and failed to recolonise the rest of northern Europe during
the post-glacial. At the same time, the seasonality of the
climate round the Mediterranean helps to diversify the soil
habitat, enabling many more species to co-exist.
One factor underlying patterns of diversity which is more
theoretical and harder to verify derives from the ecology of
the groups. Some generalist predators such as scorpions and
solifugids may have such broad niches that rather few
species can coexist in an area, although the regional
diversity in such groups can be high if the individual
species have small ranges, and species complementing
occurs on a smaller scale than usual.
Several recent estimates have suggested that the true
diversity of soil fauna, in common with most invertebrates,
may be ten times or more than the number of described
species (Erwin, 1982; May, 1988).
1. Biological Diversity
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This is a condensed version of a consultancy report
prepared by B.C. Eversham, A.S. Jolliffe and B.N.K. Davis
of Monks Wood Experimental Station, a unit of the Institute
of Terrestrial Ecology (Natural Environment Research
Council). The full unpublished report is entitled: Soil fauna
biodiversity: a preliminary global review. Project
T13061A1.
1. Biological Diversity
12. FISHES
THE DIVERSITY OF FISHES
Fishes make up the most abundant class of vertebrates, both
in terms of numbers of species and of individuals. They
exhibit enormous diversity in size, shape, biology, and in
the habitats they occupy. They are also the least known of
vertebrates. It is clear, however, that the group of animals
popularly termed fishes is defined by the retention of
primitive vertebrate features (aquatic, gills, fins, ’cold-
blooded’) and the extant groups include several rather
distantly-related evolutionary lineages. The first jawed
vertebrates, around 500 million years ago, were fishes, and
the first tetrapod land vertebrates arose from among the
fishes around 400 million years ago.
There are in excess of 22,000 described species of fish.
Vertebrates as a whole comprise around 43,000 species;
thus, approximately half of all described vertebrates are
fishes. Given that some 200 new species of fish have been
described annually in recent years, probably well over half
of all vertebrate species are fishes.
The great majority is comprised of bony fishes, mainly
teleosts (advanced jawed fishes); in addition, there are
around 800 species of cartilaginous fish (sharks, rays,
chimaeras) and 70 jawless fish (lampreys and hagfishes).
Fishes range in size from around lcm (as shown by a
Philippines Goby Pandaka pygmaea, which is about 1.2cm
in adult length, and another in the Indian Ocean, about
lcm) to the Whale Shark Rhincodon typus, which attains
15m. Some fish, typified by eels, are long and slender,
others are globular; some are almost colourless, others are
brilliantly coloured; some are fast and graceful, others
sedentary.
They occupy almost every kind of aquatic habitat, ranging
from sub-zero waters under the Antarctic icecap to near-
boiling hot springs, and in water that is almost pure or
highly saline. Many occupy the lightless ocean depths, a
few dozen inhabit lightless cave systems (and some have
lost both eyes and skin pigment).
Liquid water in lakes and rivers totals around 126,000km’,
equivalent to 0.0093 % of the total volume of liquid water
in the world. The oceans comprise about
1,320,000,000km?, or 97% of the total. More than 8,400
fish species, or about 40% of all fishes, live in freshwater.
There is thus around 100,000 km? of water for each marine
species but a mere 15km’ for each freshwater species: a
difference of several orders of magnitude.
It has been calculated that some pelagic marine species may
attain population levels of 10'* individuals, although a more
typical value might be 10°. The mean value for freshwater
species has been estimated to range down to 10°. Given the
different water volume available per species, this represents
a possible ten-fold decrease in water volume per individual
in freshwater over marine species. This is not inconsistent
with the greater net primary productivity per unit area, and
greater plant biomass, in freshwater as compared with
marine habitats.
116
Fishes provide the major world source of food derived from
wild animals. Whether assessed in terms of tonnage traded
or proportion of total dietary protein, fishes are a global
resource of the first magnitude. Although the tropics
generally have far higher species richness and endemism
than temperate or arctic regions, and include 50% and 30%
of the world’s open water and continental shelf water,
respectively, tropical fisheries contribute only about 16% of
world fish production (Longhurst and Pauly, 1987).
Table 12.2, modified from Nelson (1984), lists the orders
of extant fishes. Most orders are geographically very
widespread, with representatives in the Atlantic, Indian and
Pacific Oceans and/or on most continents: those with less
wide distributions are noted in the table. Also listed are the
numbers of families, genera and species in each order, with
estimates of the number of species in marine and freshwater
habitats.
We have made no attempt to deal comprehensively with the
biodiversity of fishes, but have concentrated on aspects of
species diversity, and include below material dealing with
species richness and endemism in freshwaters, and notes on
subterranean and coral reef fishes.
FRESHWATER FISHES: SPECIES RICHNESS AND
ENDEMISM
Estimates have been made of species richness on major
landmasses (Table 12.1), and detailed information is now
available for a few families, but Tables 12.6 to 12.10 below
are a first preliminary attempt to collate data on species
richness and endemism of indigenous freshwater fishes on
a global scale. Summary data on rivers and lakes are
represented graphically in Figs 12.2 and 12.3.
Table 12.1 Freshwater fishes: species
richness by continents
South America 2200
Africa 1800
Asia 1500 '
North America 950
Central America 354
Europe 250
Australia 170
New Zealand 27.2
Sources: Estimates cited in Nelson, J.S. 1984. Fishes of the World,
2nd edn. John Wiley and Son, New York.
Notes: ' Estimate probably should be much higher, (Nelson, 1984).
? Mostly diadromous.
In contrast to practice in other parts of this book, data on
species diversity are presented in terms of water bodies
rather than country units. River systems, for example,
frequently cross several country boundaries or themselves
constitute the boundary, making a country approach to data
compilation more difficult and biologically less meaningful.
Introduced species are excluded from the counts wherever
possible, as are subspecies (although some information
sources are too imprecise to allow this in all cases; these
Fishes
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exceptions are recorded in the notes). In general, river
systems or lakes were included only when estimates of both
the total and the endemic fish fauna were available. Several
very important rivers for which no useful data could be
traced (e.g. the Ganges, Irrawaddy, Sikiang, etc.) have
been excluded. For some of these, information does exist,
but is too outdated or incomplete to use.
Faunal knowledge
The quality and extent of faunal studies vary widely from
country to country. The geography of countries, which are
artificial constructs, often bears little relationship to the
geography of water bodies, which are natural. Because the
great majority of faunal inventories are on a sub-national
basis, substantial gaps in the coverage of a multinational
river system frequently result. The probability of such
incompleteness should be borne in mind when using the
figures given below. In general, data quality is highest in
developed countries, where species richness is lowest, and
faunal inventory will probably remain least satisfactory in
systems that cross several developing country boundaries
(e.g. the Mekong), where species richness is undoubtedly
high. It is of some concern that ichthyologists do not know
with precision how many species of fish exist in
multinational rivers, and often have but sparse knowledge
of the fauna of rivers where fishes are an important human
food resource. The ability to provide appropriate
Management remains correspondingly impoverished.
Taxonomic knowledge
Difficulties concerned with the taxonomic status of fishes
arise from scientific disagreement or ignorance. Freshwater
fish species vary in morphology throughout their range as
a result of genetic, dietary and other factors. A specimen
from one region may therefore have received a different
taxonomic name from specimens from another population
of the same species in another region. This result of
parochial taxonomic research can be corrected when a
taxonomist has access to a suitably large sample on which
to work. A similar problem occurs when different
taxonomic status is given to the same species by different
authors so that the same biological species can appear under
different names in different faunal lists. When uncritical
overviews have been undertaken the same species can
appear as two or more nominal species in the same list,
thereby artificially inflating the number of species. Without
the time to compare specimens and refer to the original
descriptions, the number of species (both widespread and
endemic) quoted in this document is based on a reasonable
interpretation of the published literature and reference to
people actively working in particular fields.
What are ‘freshwater fishes’?
Freshwater fishes are customarily categorised as primary or
secondary. This categorisation is essentially ecological not
taxonomic, although based on families. Primary freshwater
fishes, in this usage, are those families with little salt
tolerance (stenohaline) and therefore confined to fresh
waters. This has meant that the sea is a barrier and their
current distribution is a result of physiographical events.
The families Cyprinidae, Characidae and Cobitidae are
119
Fishes
examples of this group. Secondary freshwater fish families
contain species which mostly live in fresh water but have
some degree of salt tolerance and can cross salt waters. The
Cichlidae, for example, are in this category. There is an
accepted third category, the peripheral fishes, containing
families that do not conform to either of the other two
categories. Some may spend most of their life in fresh
waters; others live in brackish waters. Marine families with
representatives in fresh waters are also grouped here along
with some anadromous or catadromous fish. The usefulness
of categorising fish on the basis of salt tolerance had been
challenged by Rosen (1974) who thought that fish should be
regarded as continental or oceanic.
A pragmatic approach has been taken to determining which
species to include in these estimates. In general, if most
members of the species live in the sea, isolated freshwater
populations are not included below. Some arbitrary
decisions have been made. Anguillid eels have been
excluded on the grounds that they breed in the sea, can
move overland, and no populations isolated in fresh water
are known.
Endemism
In normal biological usage, an endemic species is one
confined to some given area, which may be defined as a
site, a country, a continent or, in this compilation, a
discrete river system or lake. If, however, a list of fish is
constructed on the basis of river systems the full picture of
very localised species will not emerge. For example, most
riverine endemic fish species live in head-waters and often
in very short stretches of river. The geophysical process of
head-water capture has frequently resulted in one highly
localised species living in two or more much larger river
systems yet, in reality, being confined to a world
distribution of just a few square miles. It has not been
possible for the present document to compile adequate data
on species which are local or regional endemics but are not
endemic to one particular river system. They are not
included below and it should be borne in mind that there
are many more species of fishes with extremely localised
distributions than is apparent here.
Species richness in rivers and lakes
The number of fish species present in subtropical and
tropical rivers is highly correlated with the area of the river
basin; temperate rivers show a similar pattern although the
number of species rises more steeply with increasing basin
area in tropical systems than in higher latitudes
(Welcomme, 1979). The relationship appears to break down
at high latitudes, where some tundra rivers are very
extensive but have few fish species. Data gathered by
Welcomme (1990) and Daget and Economidis (1975) are
tabulated (Table 12.3) and shown graphically (Fig. 12.1).
Lake area is in general positively correlated with species
richness, but a variety of additional factors may be
involved. On a global scale, surface area and latitude
together account for about one-third of the overall variation
in species number (Barbour and Brown, 1974). For a
sample of 14 lakes in North America, these factors
accounted for most of the variation in species number, the
1. Biological Diversity
Figure 12.1
10,0
1,0
4
Number of fish species
Number of fish species and river basin area
00 =
ie *
E @)
00 |-
= x
00
10
both axes on log scale
Latin America
4
10
100
River basin area (1000 km square)
Source: Data in Welcomme, R.L. 1990. Status of fisheries in South American rivers. Interciencia 15(6):337-345; and Daget, J. and Economidis,
P.S. 1975. Richesse spécifique de l’ichtyofaune de Macédoine orientale et de Thrace occidentale (Gréce). Bulletin du Muséum National d’Histoire
Naturelle, 3e série, no 346, écologie générale 27:81-84.
Table 12.3 Numbers of fish species and river basin area
RIVER
Latin America
Sucio
Paz
San Tiguel
Paraguay
Uruguay
Magdalena
Negro
Parnaiba
Madeira
Orinoco
Parana, La Plata
Amazon
Shire
Cavally
Sokoto
Oueme
Kafue
Ruaha
Tana
Comoe
Gambia
BASIN AREA RIVER
(km?)
794
1,884
2,985
181,970
223,872
256,622
331,131
362,000
691,831
950,000
3,100,000
5,711,000
3,981
5,012
9,441
19,953
22,387
35,481
39,811
44,668
59,566
66,834
70,795
83,176
Sassandra
Bandama
Cunene
White Nile
Senegal
Kasai
Volta
Chari
Ubangui
Niger
Zambezi
Zaire
Balkans
Aspropotamos
Laspopotamos
Loutos
Marmaras
Potamos
Bospos
Kossithnos
Kompsatos
Filiouris
Nestos
Strymon
Evros
NUMBER OF
SPECIES'
BASIN AREA
(km?)
84,140
100,000
169,824
229,087
342,768
357,273
378,443
575,440
668,344
1,100,000
1,280,000
3,968,000
129
180
211
17,035
52,788
Source: Data in Welcomme, R.L. 1990. Status of fisheries in South American rivers. Interciencia 15(6):337-345; and Daget, J. and Economidis,
P.S. 1975. Richesse spécifique de I’ichtyofaune de Macédoine orientale et de Thrace occidentale (Gréce). Bulletin du Muséum Nesoned d'Histoire
Naturelle, 3e série, no 346, écologie générale 27:81-84.
Note: * Total fish number given above will differ in some instances from numbers given in the main set of tables (12.6-12.10) because of different
original data sources.
Fishes
Table 12.4 Number of fish species, lake area and latitude
LAKE NUMBER OF SURFACE LATITUDE
SPECIES' AREA (km?)
Africa
Albert 46 5346 1.7°N
Bangweulu 68 2072 11.1°S
Chad 93 17500 13.0°N
Chilwa 13 673 15.3°S
Edward 53 2150 0.5°S
Kivu 17 2370 2.0°S
Malawi 245 28490 12.0°S
Mweru 88 4413 9.0°S
Nabugabo 24 30 0.6°S
Rudolf 37 9065 3.5°N
Rukwa 22 3302 8.0°S
Tana 18 3626 12.0°N
Tanganyika 214 32893 6.0°S
Victoria 177 69484 1.0°S
Canada
Athabasca 21 7154 59.2°N
Big Trout 24 616 53.8°N
Great Bear 12 31153 66.0°N
Great Slave 26 27195 61.4°N
Keller 13 406 63.9°N
Kootenay 19 399 49.5°N
La Ronge 19 1425 55.0°N
Opeongo 22 60 45.7°N
Great Britain
Loch Lomond 15 71 56.1°N
Windermere 9 15 54.3°N
Guatemala
Peten 23 98 17.0°N
Yzabal 48 684 15.5°N
Italy
Maggiore 21 212 46.0°N
Japan
Biwa 46 676 35.2°N
Mexico
Chapala 14 1080 20.2°N
Patzcuaro if 111 19.6°N
Zirahuén 5 8 19.4°N
Nicaragua-Costa Rica
Nicaragua 40 8264 11.5°N
Peru-Bolivia
Titicaca 18 9065 16.0°S
LAKE NUMBER OF SURFACE LATITUDE
SPECIES’ AREA (km?)
Philippines
Lanao 20 357 7.9°N
USSR
Aral Sea 17 64500 45.0°N
Baikal 50 31500 54.0°N
Balkhash 5 18500 46.0°N
Beloe 22 1125 60.2°N
Black Sea 156 423488 43.0°N
Caspian Sea 74 436 42.0°N
Gusinoe 13 165 51.2°N
Issyk Kul 11 6206 42.0°N
Ladoga 48 18400 61.0°N
Leprindo 14 24 56.5°N
Onega 28 10340 61.5°N
Pestovo 17 2 58.3°N
Sea of Azov 17 38000 46.0°N
Seliger 21 221 57.2°N
Taimyr 13 4650 74.5°N
Teletskoe 14 231 51.6°N
USA
Black 10 5 34.7°N
Canandaigua 37 41 42.8°N
Cayuga 60 171 42.7°N
Erie 113 25719 42.2°N
Huron 99 59596 44.5°N
Jones 13 1 34.7°N
Keuka 30 44 42.5°N
Michigan 114 58016 44.0°N
Ontario 112 19477 43.5°N
Otisco LiZ/ 10 42.8°N
Owasco 10 85 42.8°N
Salters 14 1 34.7°N
Seneca 39 174 42.6°N
Singletary 14 3 34.6°N
Skaneateles 14 54 42.8°N
Superior 67 82414 47.5°N
Waccamaw 36 36 34.3°N
Walnut 30 1 42.6°N
White 19 5 34.6°N
Yugoslavia-Albania
Ohrid 17 347 41.0°N
Source: Data in Barbour, C.D. and Brown, J.H. 1974. Fish species diversity in lakes. The American Naturalist 108 (962):473-489.
Note: ‘ Total fish number given above differs in some instances from numbers given in the main set of tables (12.6-12.10) because of different
original data sources.
very strong effect of latitude probably a reflection of
climatic severity and isolation from colonisation sources. In
contrast, in a sample of 14 lakes in tropical Africa, surface
area, depth and conductivity were the primary factors
involved (increasing depth in a sense represents an
increased area available to non-pelagic fishes). Select data
used by Barbour and Brown are given in Table 12.4.
SUBTERRANEAN FISHES
At least 47 species of fishes are either cave-adapted or have
cave-adapted populations. These highly localised
populations are widely distributed across the globe, from
about 38°N southward to the Tropic of Capricorn.
Information on these fishes is given here in order to
illustrate a facet of vertebrate biodiversity that is little-
121
known although of great intrinsic interest and of scientific
value in illustrating aspects of the evolutionary process.
Taxonomic and distributional data are summarised in Table
12.11 and site localities mapped in Fig. 12.4.
These 47 cave species represent seven orders and 13
families. Although frequently called cave fishes, this is not
wholly accurate as some live in honeycombed rocks
(aquafers) in which there are not necessarily any caves that
can be entered by humans. Indeed, some species are only
known from artesian wells that have penetrated these
aquafers. It is therefore better to refer to these fishes as
‘subterranean’, ‘cave-adapted’ or troglobionts.
Characteristic of such species is a marked trend toward
1. Biological Diversity
eyelessness, lack of pigment and low metabolic rate. It is
interesting to note that similar physical characteristics have
evolved in some freshwater species confined to rapids and
torrents in Africa and South America. These torrenticolous
species have presumably lost their eyes and body pigment
as a result of a lack of light in their habitat under stones
and rocks in turbid rapids.
Of the 13 families which include cave-adapted fishes, nine
are among the primary freshwater group. Indeed, the
Homalopteridae, Ictaluridae, Pimelodidae,
Trichomycteridae, Cyprinidae, Cobitidae and Amblyopsidae
have particularly narrow salinity requirements. The families
Ophidiidae, Synbranchidae and Eleotridae are primarily
marine. The subterranean members of these families live
near the coast in caves where they have been trapped in
some cases, by land uplift. In all cases, cave species form
a very small minority of the species in their respective
families.
Population sizes are generally unknown. Nemacheilus
smithi, for example, is known from just one specimen. Only
two cave-adapted forms have been bred in captivity. An
eyeless population of Astyanax fasciatus is on widespread
sale as the ‘blind cave tetra’. The blind form of this species
breeds true, yet if mated with the above ground (epigean)
form, as happens in nature, a complete range between
eyeless and fully-eyed, and depigmented and fully
pigmented, forms will result. Most laboratory based
behavioural studies have been conducted on this species
(e.g. Wilkens, 1971). Some observations on Phreatichthys
were made by Ercolini and Berti (1975).
Studies on subterranean fishes in the wild are lacking.
There is some evidence that breeding is seasonal and related
to the influx of water into the subterranean environment.
The young of Caecobarbus geertsi are only found after the
rainy season (M. Poll, pers. comm.).
Because of the conspicuous superficial differences between
a subterranean (hypogean) species and its epigean relatives
it had been considered normal practice to allocate a
hypogean species to a different genus. This action is now
considered to be phylogenetically unjustified (Roberts and
Stewart, 1976; Banister, 1984) and published nomenclatural
changes are used in the species list below.
Not all cave fishes show the same degree of non-
development of eyes or pigment. Some have very small
eyes (are microphthalmic) or have eyes covered with skin,
some are lightly pigmented. Such species can be regarded
as not yet fully cave-adapted. The acquisition of extreme
cave morphology implies the passage of time and this notion
has been used by some authors (e.g. Wilkens, 1982) to
argue that the fully cave-adapted species have been in their
environment longer than those that are partially adapted.
This argument involves the questionable assumption that
evolutionary rates are the same in all species. These
arguments also do not take account of the evidence for
neoteny in cave fishes (Gould, 1977; Banister, 1984).
The subterranean fishes are of particular scientific value in
exemplifying dramatic evolutionary phenomena. Within
seven orders and 12 families of fishes there are 46
examples of parallel evolution occurring in similar
environments. These evolutionary microcosms are often
now under threat. The waters in which these species live
and have evolved are a final sump for water soluble
chemicals used on land. In the regions where subterranean
fish live, water is often at a premium for human
consumption and tapped for that purpose (the only habitat
of Satan eurystomus is also the water supply for San
Antonio, Texas).
CORAL REEF FISHES
Coral reef fishes are those associated with coralline
structures. Many of these species can also occur in habitats
other than coral reefs and in regions outside the geographic
range of reef-building corals (Sale, 1980). Coral reefs are
tropical, shallow water ecosystems, largely restricted to the
area between the latitudes 30°N and 30°S (see Chapter 23).
These complex systems are highly productive, a result of
efficient recycling, high nutrient retention, and a structure
which provides habitat for a great range of organisms
(UNEP/IUCN 1988a,b,c).
Central parts of the Indo-West Pacific contain the highest
number of reef fish species (Ehrlich, 1975), and richness
decreases with increasing distance from this core area. Sale
(1980) considers that this general pattern cannot be
accounted for entirely by ecological hypotheses based upon
latitudinal gradients in diversity, but may be due to
historical factors. The origin and maintenance of high
diversity is subject to debate. One view is that high
diversity is sustained on reefs because of resource
partitioning between species, fish assemblages being
equilibrium communities (Dale, 1978; Robertson and
Lassig, 1980; Smith and Tyler, 1972). An opposing view
is that these communities are non-equilibrium unstable
systems, and that species abundance is determined through
independent differential responses to unpredictable
environmental changes (Sale, 1977, 1978, 1980;, 1978;
Sale and Williams, 1982).
Most reef fish species are relatively rare in terms of
individuals in the community. Thus, at Toliara (south-west
Madagascar) only about 25% (136) of the total number of
fish species present were ranked as abundant (Harmelin-
Vivien, 1989). Many families of coral reef fishes have a
circum-tropical distribution, although there are pronounced
differences at species level; the number of reef fish species
within a single zoogeographic region varies between 100s
and 1,000s. Most families in tropical seas include species
that occur in the coral reef fauna, and some families are
almost entirely restricted to reefs, such as Chaetodontidae,
Scaridae, and Labridae. Within the demersal component
(feeding on benthic organisms), the families Acanthuridae,
Balistidae, Belennidae, Holocentridae, Ostraciodontidae,
Pomacentridae (damselfish) and Serranidae tend to
dominate. Principal pelagic families associated with reefs,
other than the top predators such as Carangidae, Sphyraena
and sharks, include Atherinidae (silversides), Pomacentridae
and small lutjanids such as Caesio and its relatives
(Longhurst and Pauly, 1987).
Small-sized species tend to predominate, although the range
is from 2-3cm for some Eviota species to over 5m for some
sharks. Fish distribution is highly heterogeneous within a
particular geomorphological reef zone because of stochastic
processes involved in fish larvae settlement (Gladfelter er
al., 1980). Complexity in reef structure contributes to
species richness among reef fish by providing a wider
variety of niches. On a local scale, fish community
structure varies markedly between reef flat and outer reef
slope; these zones are subject to different environmental
factors affecting egg type, size-class categories, and feeding
ecology. Other zones, including boulder tract, seagrass beds
and deep outer flagstone all harbour characteristic fish
assemblages.
There is a strong positive correlation between coral and fish
species richness at given sites, although this is less evident
on a small scale within reef zones (Table 12.5). It has also
been suggested that there is a positive correlation between
the degree of live coral cover and species richness and
abundance of reef fishes (Bell and Galzin, 1984). In
addition, the presence of dietary specialist fish species is
often related to specific coral growth forms; for example,
the exclusive coral feeders in the Chaetodontidae are
positively correlated with the abundance of tall-branched
coral colonies (Bouchon-Navarro et al., 1985).
Numbers of reef fishes and
coral species
Table 12.5
CORAL REEF SITE NUMBER OF
FISH SPECIES
NUMBER OF
CORAL SPECIES
Great Barrier Reef (Australia) 2,000 500
New Caledonia 1,000 300
French Polynesia 800 168
Heron Island (Great Barrier) 750 139
Society Islands 633 120
Toliara (Madagascar) 552 147
Aqaba 400 150
Moorea (Society Is) 280 48
St Gilles (Réunion) 258 120
Tutia Reef (Tanzania) 192 52
Tadjoura (Djibouti) 180 65
Baie Possession (Réunion) 109 54
Kuwait 85 23
Hermitage (Réunion) 81 30
Source: Data from Harmelin-Vivien, M.L. 1989. Reef fish
community structure: an Indo-Pacific comparison. In: Harmelin-Vivien,
M.L. and Bourligre, F. (Eds), Vertebrates in Complex Tropical
Systems. Springer-Verlag, New York.
References
Banister, K.E. 1984. A subterranean population of Garra bareimiae
(Teleostei: Cyprinidae) from Oman, with comments on the concept
of regressive evolution. Journal of Natural History 18:927-938.
Barbour, C.D. and Brown, J.H. 1974. Fish species diversity in lakes.
The American Naturalist 108 (962):473-489.
Bell, J.D. and Galzin, R. 1984. The influence of live coral cover on
coral reef fishes communities. Marine Ecology
Progress Series 15(3):265-274.
Bouchon-Navarro, Y., Bouchon, C., and Harmelin-Vivien, M.L.
1985. Impact of coral degradation on a chaetodontid fish
assemblage (Moorea, French Polynesia). Proceedings of Sth
International Coral Reef Symposium 5:427-432.
Daget, J. and Economidis, P.S. 1975. Richesse spécifique de
Vichtyofaune de Macédoine orientale et de Thrace occidentale
(Gréce). Bulletin du Muséum National d'Histoire Naturelle. 3e
série, no 346, écologie générale 27:81-84.
123
Fishes
Dale, G. 1978. Money-in-the bank: a model for coral reef fish
coexistence. Environmental Biology of Fishes 3(1):103-108.
Ehrlich, P.R. 1975. The population ecology of coral reef fishes.
Annual Review of Ecology and Systematics 6:211-247.
Ercolini, A. and Berti, B. 1975. Light sensitivity experiments and
morphological studies on the blind phreatic fish Phreatichthys
andruzzi Vinciguerra from Somalia. Monitore Zoologico Italiano
(NS) Suppl. 6:29-43.
Gladfelter, W.B., Ogden, J.C. and Gladfelter, E.H. 1980. Similarity
and diversity among patch reef fish communities: a comparison
between tropical western Atlantic (Virgin Islands) and tropical
central Pacific (Marshall Islands) patch reefs. Ecology 61(5):1156-
1168.
Gould, S.J. 1977. Ontogeny and Phylogeny. Belknap Press of Harvard
University Press. ix + 5O1pp.
Harmelin-Vivien, M.L. 1989. Reef fish community structure: an Indo-
Pacific comparison. In: Harmelin-Vivien, M.L. and Bourliére, F.
(Eds), Vertebrates in Complex Tropical Systems. Springer-Verlag,
New York. Pp.21-60.
Longhurst, A.R. and Pauly, D. 1987. Ecology of Tropical Oceans.
Academic Press Inc., San Diego, London.
Nelson, J.S. 1984. Fishes of the World, 2nd edn. John Wiley and Son,
New York.
Roberts, T.R. and Stewart, DJ. 1976. An ecological and systematic
survey of fishes in the rapids of the lower Zaire or Congo river.
Bulletin of the Museum of Comparative Zoology 147(6):239-317.
Robertson, D.R. and Lassig, B. 1980. Spatial distribution patterns and
coexistence of a group of territorial damselfishes from the Great
Barrier Reef. Bulletin of Marine Science 30:187-203.
Rosen, D.E. 1974. Phylogeny and zoogeography of salmoniform fishes
and relationships of Lepidogalaxias salmondroides. Bulletin of the
American Museum of Natural History 153 2:265-326.
Sale, P.F. 1977. Maintenance of high diversity in coral reef fish
communities. American Naturalist 111:337-359.
Sale, P.F. 1978. Coexistence of coral reef fishes: a lottery for living
space. Environmental Biology of Fishes 3(1): 85-102.
Sale, P.F. 1980. The ecology of fishes on coral reefs. Oceanography
and Marine Biology Annual Review 18:367-421.
Sale, P.F. and Williams, D.Mc.B. 1982. Community structure of coral
reef fishes: are the patterns more than those expected by chance?
American Naturalist 120:121-127.
Smith, C.L. and Tyler, J.C. 1972. Space resource sharing in a coral
reef fish community. Bulletin of the Natural History Museum Los
Angeles Science Bulletin 14:125-170.
UNEP/IUCN 1988a. Coral Reefs of the World. Volume 1: Atlantic and
Eastern Pacific. UNEP Regional Seas Directories and
Bibliographies. TUCN, Gland, Switzerland and Cambridge,
UK/UNEP, Nairobi, Kenya. 373pp., 38 maps.
UNEP/IUCN 1988b. Coral Reefs of the World. Volume 2: Indian
Ocean, Red Sea and Gulf. UNEP Regional Seas Directories and
Bibliographies. TUCN, Gland, Switzerland and Cambridge,
UK/UNEP, Nairobi, Kenya. 389pp., 36 maps.
UNEP/IUCN 1988c. Coral Reefs of the World. Volume 3: Central and
Western Pacific. UNEP Regional Seas Directories and
Bibliographies. TUCN, Gland, Switzerland and Cambridge,
UK/UNEP, Nairobi, Kenya. 329pp., 30 maps.
Welcomme, R.L. 1979. Fisheries ecology of floodplain rivers.
Longman, London and New York.
Welcomme, R.L. 1990. Status of fisheries in South American rivers.
Interciencia 15(6):337-345.
Wilkens, H. 1971. Genetic interpretation of regressive evolutionary
processes: studies on hybrid eyes of two Astyanax cave populations
(Characidae, Pisces). Evolution 25:530-544.
Wilkens, H. 1982. Regressive evolution and phylogenetic age: the
history of the colonization of freshwater of Yucatan by fish and
Crustacea. Bulletin of the Association of Mexican Cave Studies
8:237-244 and Bulletin of the Texas Memorial Museum 28:237-244.
Chapter abridged from a consultancy report by Keith
Banister, with additional material by WCMC staff (general
introduction, reef fishes).
1. Biological Diversity
Table 12.6 Freshwater Fishes: Eurasia
Baltic Sea basin
Neva
Dvina
Vistula
Black Sea basin
Danube
Dniepr
Dniestr
Kuban
Don
Crimea
Sakayra basin (Turkey)
Caspian Sea basin
Volga
Ural
Terek
Kura & Araxes
Sefid & Atrek
Aral Sea basin
Amu-Darya
Syr Darya
Issy-Kul Lake basin
Lake Balkash basin
Tarim basin '
White Sea drainage
N Dvina
Pechora
Arctic Ocean basin
Ob
Yenisei (excl. Lake Baikal)
Lake Baikal
Lena
Kolmya
Bering Sea drainage
Anadyr
Kamchatka
Pacific Ocean drainage
Amur
Yalu
Hong Ha (Red River may
include brackish water species)
North Vietnam rivers
South Vietnam rivers 2
Mekong
Malayan Peninsula ?
Tasek Bera swamp (Malaysia)
Gombak (Malaysia)
Japan
Overall *
No of
species
¢120
No of
endemics
ooo
oooooouw
oo
_
6?
c4
77
ov
very few
?
3 -83
c 53
No of No of
species endemics
Lake Biwa (Japan) © < 63 8
Philippines ‘
Lake Lanao (Mindanao) ® c 24 c18
Indonesia
Kapuas (Kalimantan) ° c 250 ¢35
Java ec 100 cé6
Lake Poso (Sulawesi) ® 10 8
Papua New Guinea
Fly river 7 103 17
Northern rivers c 84 c 36
Sri Lanka #
54 <7?
Indian Ocean drainages
Mae Khong '° 215 15
Lake Indawgyi 43 2
Lake Lortak 13 ie)
Lake Inlé 28 7-8
Nepal (rivers Arun, Trisuli 101 Oo
Mardi-Kola, all Ganges head-waters)
Indus 147 22
(Kabul, Chamkani-Kurram, 45 10?
Zhob Gowmal, all Indus
head-waters)
Tigris & Euphrates 62 5?
Endorheic basins: Mongolia
Ugiy Nuur 7 ie)
Biger Nuur 8 0
Boon Tsagaan Nuur 2 fe)
Endorheic basins: China
Upland lakes of Yunnan "' 65 44
Er-Hai (Yunnan) '? 6 5
Endorheic basins: Afghanistan-Iran
Helmand-Sistan basin 27 1-5?
Hari-Tedzhen 12 fe)
Murgals 15 ie}
Lake Reza lyeh (Urmin) 14 5
Arabian Peninsula and Levant
Oman mountains 3 2
Red Sea, Gulf of Aden 8 7
and Wadi Hadramut systems
Rub al Khali drainage 3 3
Jordan river drainage 24 12
Azraq Oasis “* 1 1
Western and southern Europe
(excluding the river basins considered elsewhere)
Lake Ohrid '® 17 3
Europe '® 76 10
The rivers are arranged roughly clockwise from the Baltic Sea
drainage. Major lakes are included along with their tributaries, as most
literature treats the ichthyofauna on a regional or basin basis. Within
an Eurasian context this treatment is biologically rational, as many of
the lake fishes are anadromous.
Regrettably, no reliable data could be found for several major river
systems in this region; these include the Hwang Ho, Sikiang,
Irrawaddy, Ganges, and rivers of peninsular India (notably the
Godaveri, Cauvery and Narmada).
er Awe
Fishes
Notes
In toto there are 16 species endemic to the Aral Sea basin.
Of these species, about 80 are brackish water inhabitants or largely marine.
The freshwater fishes of Peninsular Malaysia are divided into 3 faunal zones: the northwest, the northeast and central, and the south zone. The
numbers of freshwater fishes in each division are given respectively as 47, 98 and 58. The number of species listed in various articles as endemic
varies extremely widely. Some reliance can be placed on the total number of primary freshwater fish, at least as to the order of magnitude, but
very little on the number of endemic species.
This figure probably includes some euryhaline species.
The total figure includes introductions and subspecies.
The number of endemic species will be higher if the immediately adjacent rivers were included (Roberts 1989).
The endemic tally would be 47 if one or more other rivers from central-southern New Guinea were included (Roberts 1978).
This lake is famed for a reported endemic species-flock of cyprinids. However, since 1962 when alien species were introduced the indigenous
fauna has become extinct (see Kornfield & Carpenter 1984; Reid 1980). Furthermore, many of the original specimens collected by Herre that
led to the idea of the Lanao species flock were destroyed during the Japanese invasion in World War II. The number of species on other islands
varies widely but has not been the subject of detailed listings.
It is not known if 2 of these species still survive. Introductions are likely to be responsible for their possible extirpation (Kottelat 1990).
This figure apparently includes about 40 brackish water and introduced species. The endemics are loaches and homalopterids from head-water
streams.
These figures include subspecies as well as 2 endemic genera. It also seems that endemic in this context means very limited distribution but in
more than one water body (Li 1982).
All these species are cyprinids.
The fauna of this region, especially of Lake Urmin, is very badly in need of re-examination.
There have been very many instances in historical times of translocation of fishes within and into this area that well over half the fishes now
living in the system are not indigenous. These have not been included above.
One of the ‘endemics’ occurs in immediately adjacent lakes.
The European fish fauna is richest in the west and becomes increasingly depauperate towards the Mediterranean, Atlantic and North Sea coasts.
This trend is even more marked in the off-shore islands which were separated from continental Europe at the end of last ice age, before the full
complementof the refugia fauna had moved westwards. Only the most widespread or euryhaline forms live in Ireland, for example. The endemic
species live in Dalmatia (1), Greece (3), Spain (3), North Italy south of the Alps (1), Italian rivers draining into the northwest Adriatic (1) and
the Rhone (1). The European fishes have been much studied but rarely in a global context, and the significance of minute differences has been
given greater importance than is probably justified. Only recently has a trend started to look at European fishes in an Eurasian context, which
probably will affect the classification of the fishes quite considerably.
Table 12.7 Freshwater Fishes: North America
Total Endemic Total Endemic
species species species species
Far north Central Appalachian western drainages
Hudson Bay drainage ' 101 fo)
(Ohio system headwaters)
Ungava Bay watershed ” 18 fe) Allegheney 92 fo}
Arctic archipelago * 8 (e) Muskingum 111 (e)
(no primary freshwater fish) Monongahela 89 {e)
St Lawrence River 98 1 Little Kanawha 72 (0)
Newfoundland rivers 20 (0) Kanawha: below falls 90 fe)
(no primary freshwater fish) Kanawha: above falls 49 6
Labrador rivers 26 fo) Guyandotte 67 te)
(2 primary freshwater fish) Big Sandy 94 {e)
Northern Appalachian rivers Southeastern USA
rae!
Bec es cea ea ae fish) bit SESS ee “
Savannah 75 {e)
Central Appalachian Atlantic drainages Apalachicota drainage 86 7
Choctawhatchee 74 fo)
Edisto 55 ) Perdido 57-64 ()
Santee 90 5 Mobile Bay drainage 157 c 40
Peedee 76 1 Kissimmee river (and Lake 37 {0}
Waccamaw 51 2 Okeechobee)
Cape Fear 71 1 Suwannee (and 43 fo}
Neuse 70 ) Withlacoochee)
Tar 66 0 Mississippi-Missouri ° c 260 c72
fore = B Rio Grande Basin 121 69
ames P
York 49 0 California Coastal to Oregon
epbahanock ay 2 (and internal basins)
Potomac 65 1 8
Susquehanna 61 1 Eee oy ue
Sacramento system ’” 38 6
1. Biological Diversity
Total Endemic Total Endemic
species species species species
Far north Yukon and Mackenzie basins
Peace 24 0
Death Valley system 8 6 Mackensieiniver 34 0
North central basins 4 2 P
Lahontan basin 13 5 Yukon river 33 0
Bonneville basin 19 8 Lakes
Oregon lakes 15 3 Superior 44 0
Klamath river 28 6 Erie ? 99 0
N California - Oregon rivers 29 3 Ontario 95 0
Cascadia Michigan” 78-130 0
(the Columbia system north to Stikine) Huron 86 0
Columbia 45 13 Pontachartrain 716 1
Fraser 39 fo) Lahontan - see Lahontan basin
Skeena 32 io} Tahoe - see Oregon lakes
Nass 27 + Great Slave lake 36 0
Stikine 27 fe)
and tributaries
Notes
' The many recent introductions are excluded here. The drainage covers a wide range of climate zones and most of the species are in the south
of the region and are probably recent (post-glacial) migrants.
This figure includes freshwater species with some degree of euryhalinity.
Fish have only occupied this area for 14,000 years. Much of it is ice-covered in winter.
The unique endemic is a rare anadromous coregonid found only in the fresh waters of the southern tip of Nova Scotia.
Of these endemics 56 come from the Cumberland, Tennessee and Arkansas drainages, ie, a very small part of the system.
The number of species and endemic species could change substantially at any time as there is disagreement about the specific or subspecific status
of some forms, as well as known problems with hybridization.
One of the Sacramento endemics has been widely introduced elsewhere and the number of endemics would have been much higher if small,
adjacent, but quite separate rivers had been included here.
Migratory forms are included in the Yukon figures.
One endemic subspecies now extinct.
Larger figure includes tributaries.
A uw eww
Table 12.8 Freshwater Fishes: Central and South America
Total Endemic Total Endemic
species species species species
Mexico Nicaragua
Santiago 17 2 Lake Nicaragua 2 c39 (o)
forma 35 15 Pacific slopes 12 co)
Morelia 14 1 Atlantic slopes 32 {¢)
Patzcuaro 10 2 Cexeuulcs = e
irahuén 8 0 Lake Managua 26 1
San Juanico 6 2 South America
Valle de Mexico ' 5 1
Puebla plateau 4 2 Trinidad 36 5
Atonilco 7 {e) Magdalena 166 ?
Ameca 20 7 Maracaibo 108 31
Magdalena 8 1 Caribe 48 6
Armeria 11 3 Lago de Valencia 35 4
Coahuayana 9 3 Orinoco * 318 88
Balsas 27 8 Amazon ‘ c 2000 c 1800
Papagayo 4 (0) Rio Negro (Amazon) 436 35
Varde Atoyac 9 1 Lake Titicaca 20 14
Panuco 75 22 Trans-Andean region 390 ec 100
Gulf coast 21 1 La Plata ® c 550 ¢110
Papaloapan 57 9 Uruguay ec 160 e35
Notes
‘ In this region 3 additional former endemic species have recently become extinct.
The zero for the number of endemic species in the lake does not reflect the fact that it contains species of extremely limited distribution which
variously occur in associated water bodies.
So far as can be ascertained, these figures include subspecies and probably also include some not strictly freshwater fish.
This figure is extremely imprecise. Most published figures vary widely, and the relevance of detailed studies at one locality to the fauna of the
appropriate part of the subsystem is in doubt.
The total number of species in this river includes an unknown number of euryhaline species. The Parana, above the Guayra falls, has a high
proportion of endemics in its fauna which is depauperate when compared to the rest of the system.
The entities listed in the table include rivers, lakes, and one island (Trinidad).
2
126
Fishes
Table 12.9 Freshwater Fishes: Australia and New Zealand
Total Endemic Total Endemic
species species species species
Australia ' New Zealand ”
3 5 c 30 27
c110 c 105
tes
No
1
It is very difficult to categorize the Australian fishes in the same way as in other parts of the world. Strictly speaking, primary freshwater fish
number just 3, of which 2 are endemic. The total number of species living all or the major part of their lives in fresh water is about 150. Of
these, about 110 seem to be confined to fresh waters, even if they are capable of living in sea water. A further difficulty is that many of the
‘fresh’ waters are remarkably saline, especially in the desert regions. The great majority of fishes are confined to the short, peripheral, coastal
rivers. All the 110 or so species had marine ancestors and many have marine close relatives; they are either physically confined to non-marine
waters or are supposed to inhabit and breed in the freshwater parts of rivers. However, this figure could easily vary by 15% either way.
Similar problems occur in evaluating the status of New Zealand fishes, except that there are no primary freshwater fishes there.
Table 12.10 Freshwater Fishes: Africa
Total Endemic Total Endemic
species species species species
Atlantic drainage Internal drainage rivers
Senegal ' 83 3 Omo (Lake Turkana 20 1
Garbis 79 0 Chari (Chad) ’” c 162 c 25?
Tominé 36 1 Malagarazi (Tanganyika) ® >14 1
Koukouré 344 5 Ruzizi (Tanganyika) ° 92 29
Great Scarcie (Kolenté) 23 {e) Cubango (Okavango) "®
Sassandra 65 2 Natural lakes
ae ah ue Afrera (Guilietti) 2 1
Volta 2 132 8 ial ae oh
Rane 39 0 Bangwelu at 86 {0}
Gasmé 62 1 Barombi-Mbo '? 17 12
Niger 149 13 Chad "° 93 1-30
Pinas Meme 27 5 Chilwa '* 13-18 1
Rio Muni 81 36 Edward-George c55 e35
Zaire ° c 700 c 500 ves complex 1° : u
Cunene * 55 2? pie a al <
Orange-Vaal 16 5 Kiva aT UES Set
Malawi >250 >230
Cape drainage ce 338
c 1000
Rivers of the great 13 10 Mweru "8 85 fo)
escarpment and eastern plateau ® Nabugabo '® 24 5
; 20
Indian Ocean Drainage eee endMagadi 2 br }
Piitanteiriver 10 5 Tanganyika ”" >250 >230
Limpopo 49 2 Tsana (Tana) 72 c 20 1?
Zambezi 122 ce 25 Tumba * > 100 1
Great Ruaha ® >36 3 Turkana (Rudolf) 7 48 10
Tana >2 Upemba lakes 7° c 130 1
Victoria (including >250 >225
Mediterranean drainage Kyoga) 7°
; Zwai ”’ <20 3
Nile 115 26 Madagascar 7
Tunisian rivers 6 1
c 40 38
Notes
1
2
3
The 3 endemic species are only found in small headwater streams.
There is a large number of small rivers draining south from the Guinea highlands which hold many species restricted to several rivers in that
region. In the original species descriptions the localities are given but cannot be put into context as the total fauna of these rivers has not been
described. The high level of regional endemicity is not reflected in this table. Only the larger rivers (Tominé to Volta) have been studied in
sufficient detail to make an adequately reliable entry.
The Zaire figures are a consensus of the most recent estimates. Over the last few years reduction in the number of nominal species by
synonymisation has roughly equalled the descriptions of new taxa. The given figure has been based on the collections made at relatively few sites
within the vast river network. (See, in particular, Banister, 1986: 215-216.)
Comments made in Note ? apply equally to the small rivers of Angola between the Quanza and the Cunene. For example, Ansorge, made a
collection of fish in the early years of this century close to Lucalla railway station on the Lucalla river. His collection contained 25 species, of
which 11 were unique to that site. No more recent records of fish collections from that region have been located.
127
Biological Diversity
There is a very high level of endemicity in this localized Cape fauna. The main named rivers are the Berg, Breder and Buffalo rivers. The
indigenous fauna is not speciose but now there are many introductions, to the detriment of the local fauna.
This figure is based on a pre-impoundment survey in just one part of the Rufigi system.
Although 25 is the most commonly cited number of endemics in the Chari-Logone system, it seems likely that, at best, many are sub-species.
The basin fauna consists largely of widespread Nilotic fishes with a contribution of Niger-Benue faunal elements. The Chari-Benue watershed
is extremely low and the systems connect during periods of heavy rain.
The Malagarazi is a swampy river flowing sluggishly westward across a plain to Lake Tanganyika. Its poorly known fauna is Zairean in origin
as the present Malagarazi is a now isolated former part of the Zaire system.
The Ruzizi is the main inflow to Lake Tanganyika, yet it is only about 12,500 years old. At that time the water level in Lake Kivu rose to such
an extent that it overflowed southwards and the Ruzizi was formed. The upper and lower reaches of the river have different faunas and different
hydrological conditions: Upper reach - Total 27. Endemic 7; Lower reach - Total 65, Endemic 20; Common to both - Total 13, Endemic 6.
There are difficulties in evaluating the fauna of the lower reaches because of fish movements between the Ruzizi and the lake. The lake cichlids,
however, rarely penetrate far up the river. In addition to the species enumerated above, there are 3 endemic species in streams flowing from
the west into Lake Tanganyika. The streams are not meaningfully named.
The fauna of this endorheic river is essentially that of the Zambezi (q.v.). However, its upper reaches and headwaters are very poorly known.
The 86 species include those that live in the surrounding interconnecting small lakes, creeks and marshes. Lake Bangweulu does not have clearly
defined limits. Poll (1957) stated that 17 species live in the main lake.
The total number includes the species that inhabit the feeder streams and may occur in the lake itself at the feeder inflows. Of the 12 endemics,
11 are cichlids.
Chad is a rapidly dissociating lake in a shallow basin. Formerly it was much more extensive. The total number of species is that of the entire
basin. Only one species is endemic to the nucleus of the lake, but 25-30 are endemic to the entire basin.
This lake periodically dries up. The fish take refuge in residual pools or in feeder streams when this happens or when the conductivity gets too
high. The higher figure for the total number of species includes those that normally live in the feeder streams, but all must live together at times
of desiccation.
This is a series of 4 small lakes on the Tanzanian shield, Eyasi, Kitangiri, Manyara and Singida, that are the remnants of a former, much larger
shallow lake.
In the earlier literature, no distinction was made between an occurrence in a feeder stream and in the lake itself. The figures given above are,
respectively, for the lake basin and the lake, but at least one of the basin species occurs in the lake but only at the mouth of feeder streams.
Lake Malawi illustrates the uncertainties involved in compilation of this list. The three lines of species numbers above demonstrate the difference
between published figures, current knowledge, and a probable future number when the lake fauna is well known. The top line is the published
estimate. The second line is the current number of species described or known to be in press and to be published within the next year or so. The
third estimate is based on information from Prof J. Stauffer (Pennsylvania State University): "Additionally, there are at least 200 entities which
most authorities working in the lake recognize as valid species, but for which no species descriptions exist. Many of these are known by common
names in the aquarium trade. Based on the number of undescribed species which occur in the trawl samples and the fact that little is known about
the fishes inhabiting the Mozambique coast, I estimate that there are at least 1000 species which inhabit Lake Malawi. Approximately 95% of
the total fish fauna is endemic to the lake." (in litt. to K. Banister 25 June 1991.) Whichever number is most correct, only 38 species are not
cichlids.
This lake lies on a shallow watershed between the Zambezi and Zaire systems and contains fish from both systems.
The formation of this lake, an offshoot of Lake Victoria, has been dated at 4,000 years BP. All the endemics are cichlids.
These are relict, highly alkaline lakes, formerly parts of a larger lake.
There is a much higher percentage of non-cichlid endemics than in the other rift valley lakes and a much higher number of families with endemic
representatives.
The alleged one endemic is the loach Nemacheilus abyssinicus. There is considerable suspicion that the unique specimen was accidentally
translocated from a collection of middle eastern fishes into the Degen collection of fish from Lake Tsana and inadvertently described by
Boulenger (1902) as indigenous to that lake.
Although definable as a lake, it is a zone of permanent inundation up to 10 metres deep.
Of the 48 species, 36 are exclusively in the lake. The other 12 occur only in the Omo River inflow.
The Upemba lakes lie in the Kamalondo depression and are a shifting series of permanent, shallow, eutrophic lakes that are in varying contact
with the Lualaba river. Of necessity, the number of the species has to include those also present in the Lualaba.
Giving a reliable number of Lake Victoria species is very difficult as two contradictory factors are involved. First, there are an unknown number
of yet undescribed cichlid species in museum collections. Second, the recent introduction of the predatory Nile perch (Lates niloticus) into the
lake has apparently caused the extirpation of some species. The fauna of Lake Victoria is in a state of flux and the figures must be treated
correspondingly.
77 Unusually, none of the endemics are cichlids. They are cyprinids and probably spend some time in the lake as well as in the Maki river. The
lake is drying out and there is no recent information on the fish fauna.
2® All are ‘secondary’ freshwater fishes; see Introduction.
References 67.
Li Shusen 1982. Fish*fauna and its differentiation in the upland lakes
Banister, K.E. 1986. Fish of the Zaire system. In: Davies, B.R. and of Yunnan. Acta Zoologica Sinica 28(2):169-176 [In Chinese with
Walker, K.F. (Eds), The Ecology of River Systems. Dr W. Junk, English summary].
Dordrecht, Netherlands. Pp.215-224. Poll, M. 1957. Les genres des poissons d’eau douce de |’Afrique.
Boulenger, G.A. 1902. Descriptions of new fishes from the collection Annales du Musée Royal du Congo Belge. Tervuren, Sciences
made by Degen, E. in Abyssinia. Annals and Magazine of Natural zoologiques 54:1-191.
History (Series 7) 10(60):421-439. Reid, G.Mc.G. 1980. "Explosive speciation” of carps in Lake Lanao
Kornfield, I. and Carpenter, K.E. 1984. Cyprinids of Lake Lanao, (Philippines) - fact or fancy? Systematic Zoology 29:314-316.
Philippines: taxonomic validity, evolutionary rates and speciation Roberts, T.R. 1978. An ichthyological survey of the Fly river in Papua
scenarios. In: Echelle, A.A. and Kornfield, I. (Eds), Evolution of New Guinea with descriptions of new species. Smithsonian
Species Flocks. University of Maine at Orono Press, Orono. Contributions to Zoology 281:1-72.
Pp.69-84. Roberts, T.R. 1989. The freshwater fishes of western Borneo
Kottelat, M. 1990. Synopsis of the endangered Buntingi (Osteichthyes: (Kalimantan Barat, Indonesia). California Academy of Sciences
Adrianichthyidae and Oryziidae) of Lake Poso, Central Sulawesi, 14:1-120.
Indonesia, with a new reproductive guild and descriptions of three
new species. Ichthyological Exploration in Fresh Waters 1(1):49- Tables prepared for WCMC by Keith Banister.
128
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135
1. Biological Diversity
13. HIGHER VERTEBRATES
The vertebrates, with around 43,000 known species, make
up a very minor proportion of the global total of some 1.7
million described species but, by virtue of their size,
adaptions and ecological role, often exert a major effect on
the structure of communities and habitats.
Vertebrates (Craniata or Vertebrata) make up the principal
sub-phylum of the three included in the phylum Chordata.
The vertebrates share the common characteristic of a hard
endoskeleton (interior skeleton) with a backbone. There are
seven Classes (Parker, 1982), of which three, namely the
Cephalaspidomorphi or Agnatha (lampreys and hagfishes),
the Chondrichythes (sharks, rays, and skates) and
Osteichthyes or Teleostomi (the bony fishes) are commonly
referred to collectively as fishes. They are the extant
members of diverse early vertebrate lineages, no longer
recognised as a monophyletic taxonomic group. Fishes
comprise nearly half of all extant vertebrate species. Data
on species richness and endemism in freshwater fishes are
presented in Chapter 12. The remaining four classes -
Amphibia (amphibians), Reptilia (reptiles), Aves (birds) and
Mammalia (mammals) - are often referred to as the higher
vertebrates or tetrapods.
THE GROUPS OF HIGHER VERTEBRATES
Amphibians
No single characteristic uniquely defines the amphibians.
All are ectotherms, using external environmental sources of
energy to regulate body temperature, and have highly
permeable skin and pedicellate teeth. Most, but not all,
have a dual life-cycle, being aquatic as larvae and terrestrial
or semi-terrestrial as adults. Amphibians are the only four-
limbed animals in which metamorphosis, the abrupt
transformation from larvae to adult, occurs.
There are currently in excess of 4,000 described species of
amphibian (Frost 1983), divided into approximately 400
genera, 34 families and three orders (Halliday er al., 1986).
The three living orders are the Urodela or Caudata
(salamanders, newts and their allies); the Anura (frogs and
toads); and the Gymnophiona (the caecilians). Amphibians
are ecologically less versatile than other higher vertebrate
groups, in general being dependent on adequately high
temperatures, moist conditions, and the availability of water
for breeding and larval development. ;
Reptiles
The most obvious feature of reptiles is their covering of
dry, horny scales, formed by localised thickenings of the
keratin layer of the epidermis. Other characteristics include
air-breathing, ectothermy - the dependence on external
sources of heat to maintain a rather variable blood
temperature - simple unspecialised ‘homodont’ teeth, and
reproduction (normally on land) via the production of
shelled eggs or live young.
Approximately 6,550 species of living reptile have been
described, classified into about 905 genera, 48 families and
136
four orders (Halliday et al., 1986). These comprise the
Chelonia (tortoises, turtles and terrapins), the Crocodylia
(crocodiles and alligators), the Rhynchocephalia (which
contains two species of tuatara Sphenodon), and the
Squamata. This last group is divided into three suborders:
Sauria (lizards); Serpentes (snakes) and Amphisbaenia
(worm lizards) (many taxonomists recognise each of these
as an order, rather than suborder). Unlike most amphibians,
most reptiles are truly terrestrial and do not require
environments rich in water; a number of species have also
been able to adapt to marine habitats. The most species-
rich, abundant and widely distributed reptile groups are the
lizards and snakes.
Birds
All birds are, like mammals, endotherms, but their
distinctive characteristic is that they possess feathers.
Feathers are an evolutionary modification of reptilian scales
which initially probably simply served a thermoregulatory
function but now, in conjunction with the development of
the forelimbs into wings, allow most birds the power of
flight. Birds have lost all teeth from their bills, and
reproduce by laying hard-shelled eggs.
Sibley and Monroe (1990) recognise 9,672 species of bird,
organised into 2,057 genera, 144 families, and 23 orders.
Birds are therefore the most diverse terrestrial vertebrate
group. They have adapted to all the major habitats of the
world, including equatorial forests, hot deserts, and the
high Arctic and Antarctic.
Mammals
Mammals are animals whose bodies are insulated by hair
(often in the form of a thick pelt or fur), which nurse their
infants with milk produced from mammary glands, and
which share a unique jaw articulation between the dentary
(the main bone of the lower jaw) and the squamosal bone of
the skull. Present-day mammals are ‘heterodont’, i.e. their
teeth are specialised to fulfil different functions, and
endothermic, i.e. their internal body temperatures are
maintained by energy generated from metabolic processes
within the body.
Corbet and Hill (1991) list 4,327 recognised mammal
species, arranged into approximately 1,000 genera, 135
families, 18 orders and two subclasses (Macdonald, 1984).
The division into subclasses reflects a separation which
occurred almost 200 million years ago between the egg-
laying Prototheria (the only survivors of which are three
Monotremes: the platypus and two echidnas) and the Theria
which bear live young. The live-bearing mammals diverged
around 90 million years ago into the groups now recognised
as marsupials (infraclass Metatheria) and the placental
mammals (infraclass Eutheria). As a class, mammals are
extremely versatile and have adapted to almost all terrestrial
and aquatic habitats. Monotremes are only found in
Australasia and marsupials are confined mainly to
Australasia and the Neotropics, but placental mammals have
spread throughout the globe, including the polar regions.
THE DISTRIBUTION OF HIGHER VERTEBRATES
Patterns of higher vertebrate distribution
As with many other organisms, species richness of land
vertebrates tends to increase at lower latitudes. Amphibians,
for example, are generally absent at very high latitudes
(although one salamander species, Hynobius keyserlingii,
ranges as far north as the Arctic circle 66.5°N) and species
richness in most groups increases progressively towards the
equator. Trends along moisture and altitudinal gradients are
superimposed upon the latitudinal trend. For example, in
North America, the greatest numbers of species are found
in areas of high rainfall, principally in the south-eastern
USA and secondarily in the north-west. Amphibian species
diversity generally declines with altitude, so that a transect
along the Equator from the Amazon basin to the crest of the
Andes reveals a gradual reduction from 81 species at 340m
to only four species above 3,500m (Duellman and Trueb,
1985). Reptiles are extremely sensitive to cold conditions
and species diversity is very low in very high latitudes.
Reptile species diversity increases towards the subtropics
and tropics, to which some groups, such as the Crocodylia,
are completely confined. Similarly, the diversity of birds
and mammals increases towards the Equator.
The geopolitical distribution of higher vertebrates
Table 13.1 is a new compilation of data on species richness
and endemism in vertebrates other than fishes, assessed on
a geopolitical basis. This table is intended to complement
the parallel compendium of flowering plant data (Table 8.3)
earlier in this book.
Figs 13.1-13.8 show select data from Table 13.1 in graphic
form. In this set of figures we have focused on single-
country endemic species of mammals, birds and
amphibians; the data are less complete for reptiles. We do
not yet have a full data set for total country numbers, and
here show (Fig. 13.2) the countries with most mammal
species. Figs 13.3-13.8 represent the same countries shown
in the higher plant graphs (Chapter 8).
Content and format
The table attempts to give realistic estimates of:
© the total number of species of mammals, birds, reptiles
and amphibians present in each country of the world
e the number of species in each group that is endemic to
each country.
‘Endemic’ in this context means that the species distribution
is entirely within the political boundaries of a given
country; they are single-country endemics, as opposed to
site or area endemics.
It is important to note that for the purposes of this table,
islands are included with their parent country (unless
separately listed). Thus, the Galapagos are included with
Ecuador, Hawaii with USA, the Canary Islands with Spain,
and so on. Some apparent anomalies in the estimates are a
result of this political aggregation; for example, the UK has
13 endemic birds listed, but 12 of these are from overseas
territories (Henderson, Inaccessible, St Helena, S Georgia
137
Higher Vertebrates
and S Sandwich Is, Tristan da Cunha). This affects the bird
data in particular for a small number of countries.
Criteria for inclusion
Certain conventions have been followed wherever possible.
e Marine cetaceans, sea turtles and sea snakes are
excluded. However, in a very few cases, especially
where data have been taken from non-primary sources,
we have been unable to establish whether cetaceans, for
example, have been included or not.
Data for birds include regular breeding species and
exclude non-breeding migrants, occasional visitors and
vagrants. It was felt that this would give a more
consistent basis for comparison, and would avoid, for
example, the problems involved in enumerating vagrants.
Data available for some countries have not allowed us to
make these exclusions and the figures will be
correspondingly inflated - for example, an estimate of
the birds of a Sahara-Sahel country will be low if only
regular breeding species are counted, but more than
twice as large if winter migrants and vagrants are
included.
Species known to be recently extirpated from or recently
introduced to a country have been excluded.
Data quality
The estimates will become increasingly accurate as more
and better data become available. Errors arise principally
because of inadequate species inventory within countries
and continual flux in the taxonomic status given to different
population groups.
Species inventory based on field survey work is to varying
degrees incomplete. Knowledge of the fauna of many
developing countries is based largely on old and
taxonomically outdated literature, often from colonial times.
Taxonomic work results in continuing changes in
nomenclature and the delimitation of species boundaries;
populations recognised by one authority as belonging to one
species will often be assigned to one or more other species
by another taxonomist.
A further complication arises from the fact that animal
distribution is dynamic not static; the geographical limits of
species change over time, either as a slow advance or
retreat of populations at the edge of a species range or as a
more rapid population collapse or colonisation event (the
latter perhaps most evident with bird populations).
We have made no systematic attempt to survey the primary
literature for taxonomic changes that post-date the published
works consulted. In general, the number of species reported
in older literature to occur in any given country will have
been both reduced by synonymy and enlarged by the
description of new species.
These factors mean that a substantial margin of error is
associated with all these data. It has not been possible to
make a rigorous assessment of the extent to which estimates
from several sources for a given parameter differ, but
informal comparisons suggest a margin of plus or minus
1. Biological Diversity
10% is quite common and greater variation is not
uncommon.
In the ‘endemic species’ columns we have attempted to
minimise problems arising from taxonomic differences by
deriving estimates for each group for almost all countries
from a single consistent source. These sources are marked
with an asterisk in the list below. In a few cases, later
estimates based on new fieldwork have been incorporated.
The ‘total species’ columns include data from a variety of
sources. These include published country or regional faunal
monographs and the WCMC species database (itself based
upon the former category of sources, but not complete for
all vertebrate classes for all countries of the world). The
extent of variety among these data sources, in terms of data
quality, publication date and place of origin, will have led
to a corresponding variety in data quality among the figures
provided.
References
* Corbet, G.B. and Hill, J.E. 1991. A World List of Mammalian
Species, 3rd edn. Natural History Museum Publications and Oxford
University Press.
138
Duellman, W.E. and Trueb, L. 1985. Biology of Amphibians.
McGraw-Hill, London, New York.
Frost, D.R. 1983. Amphibian Species of the World. A taxonomic and
geographical reference. Allen Press Inc. and the Association of
Systematics Collections, Lawrence, USA.
Halliday, T., Adler, K. and O’Toole, C. 1986 (Eds).
Encyclopaedia of Reptiles and Insects. Unwin, London, UK.
Macdonald, D. 1984 (Ed.). The Encyclopaedia of Mammals. Unwin,
London, UK.
Parker, S.P. 1982. Synopsis and Classification of Living Organisms.
McGraw-Hill, London, New York.
* Peters, J.A., Donoso-Barros, R. and Orejas-Miranda, B. 1986.
Catalogue of the Neotropical Squamata. (Part I Snakes, Part II
Lizards and Amphisbaenians). Smithsonian Institution.
Porter, K.R. 1972. Herpetology. Saunders Company, Philadelphia,
London and Toronto.
* Schwartz, A. and Henderson, R.W. 1991. Amphibians and Reptiles
of the West Indies: descriptions, distributions, and natural history.
University of Florida Press.
* Sibley, C.G. and Monroe, B.L. 1990. Distribution and Taxonomy of
Birds of the World. Yale University Press, New Haven and
London.
* Welch, K.R.G. 1982. Herpetology of Africa: a checklist and
bibliography of the orders Amphisbaenia, Sauria and Serpentes.
Robert E. Krieger Publishing, Malabar, Florida.
The
Higher Vertebrates
Table 13.1 Species richness and endemism: higher vertebrates
MAMMALS BIRDS REPTILES AMPHIBIANS
Species Endemic Species Endemic Species Endemic Species Endemic
known species known species known Species known species
ASIA
Afghanistan 123 0 456 0 103 = 6 1
Bahrain = 0 = 0 25 0 = 0
Bangladesh 109 0 354 0 119 - 19 0
Bhutan 109 0 448 0 19 = 24 0
British Indian Ocean Territory = 0 = 0 = = = oO
Brunei 155 (e) 359 () 44 - 76 tt)
Cambodia 117 0 305 te) 82 = 28 (0)
China 394 62 1100 63 282 = 190 131
Cyprus 21 0 80 2 23 1 4 0
Hong Kong 38 0 107 () 61 (e) 23 2
India 317 38 969 69 389 156 206 110
Indonesia 515 165 1519 258 511 150 270 100
Iran, Islamic Rep 140 4 - 1 164 3 11 5
Iraq 81 1 145 1 81 - 6 rt)
Israel = 2 169 ts) = = = ()
Japan 90 29 >250 20 63 28 52 35
Jordan = 0 132 (0) = = = tt)
Korea, Dem People’s Rep = 0 = 0 19 1 13 0
Korea, Rep 49 (0) - (+) 18 - 13 1
Kuwait = () 27 () 29 te) 2 (*)
Laos 173 0 481 1 66 - 37 1
Lebanon 52 () 124 0 = = = (0)
Malaysia 264 14 501 4 268 = 158 39
Maldives = 0 24 0 = = = 0
Mongolia = 6 = 0 = = = (*)
Myanmar 300 8 2867 4 203 29 75 9
Nepal 167 1 629 1 80 = 36 7
Oman 46 3 = 0 64 11 = 0
Pakistan 151 3 476 () 143 22 17 2
Philippines 166 90 395 172 193 131 63 44
Qatar - 0 - 0 17 0 = 0
Saudi Arabia = 1 59 ts) 84 5 - ts)
Singapore 57 1 118 t) = = = 0
Sri Lanka 86 12 221 20 144 75 39 19
Syria = te) 165 (e) = = = (s)
Taiwan 62 13 160 15 67 20 26 6
Thailand 251 5 616 2 298 39 107 13
Turkey 116 0 284 0 102 5 18 2
United Arab Emirates = () = 0 37 1 = 0
Viet Nam 273 5 638 12 180 = 80 26
Yemen = 1 = 8 77 25 = 1
USSR*
276 55 - 13 168 - 37 2
EUROPE
Albania 68 0 215 0 31 (0) 13 (0)
Andorra = 0 104 0 = 0 = ()
Austia 83 te) 227 te) 14 (:) 20 ()
Belgium 58 0 180 () 8 () 17 ()
Bulgaria 81 i} 242 [*} 33 0 17 i!)
Czechoslovakia 81 0 227 te) 12 0 19 0
Denmark 43 0 185 0 5 0 14 0
Faeroe Islands - () 75 0 0 ts) 0 0
Finland 60 0 230 () 5 te) 5 (:)
France 93 0 267 9 32 0 32 3
Germany 76 (0) 237 9 12 0 20 0
Greece 95 2 244 0 51 4 15 1
Hungary 72 0 203 0 15 (0) 17 (0)
Iceland 11 (°) 80 () 0 te) 0 0
Ireland 25 0 141 0 1 {0} 3 0
Italy 90 2 254 0 40 1 34 10
Liechtenstein 64 0 134 0 7 0 10 0
Luxembourg 55 0 130 0 7 0 14 0
Malta 22 () 28 (0) 8 1 1 0
Monaco = 0 = () 6 0 3 tt)
Netherlands 55 0 187 (0) 7 (0) 16 (0)
Norwa' 54 () 235 (e) 5 te) 5 0
Polan 85 0 224 0 9 0 18 0
Portugal 63 1 214 2 29 1 17 0
Romania 84 (°} 249 i*} 25 () 19 0
San Marino = te) = to) = t°) = ()
Spain 82 4 275 6 53 13 25 2
Sweden 60 0 249 0 6 0 13 0
Switzerland 75 te) 201 (0) 14 (0) 18 0
United Kingdom 50 0 219 13 8 0 iL 0
Vatican City = 0 = () = te) = 0
Yugoslavia 95 2 245 0 41 2 23 (0)
139
1. Biological Diversity
Table 13.1 Species richness and endemism: higher vertebrates (continued)
MAMMALS BIRDS REPTILES AMPHIBIANS
Species Endemic Species Endemic Species Endemic Species Endemic
known species known species known species known Species
NORTH AND CENTRAL AMERICA
Anguilla 5 to) a te) = 1 = te)
Antigua and Barbuda 7 0 = 0 9 4 2 0
Aruba = 0 - te) 10 2 1 te)
Bahamas 12 2 88 3 24 16 5 it)
Barbados 6 0 24 i*) = 3 = i)
Belize 125 0 528 0 107 2 - 0
Bermuda - ts) = 1 = = = 0
Canada 139 4 426 3 41 ie) 40 i)
Cayman Islands 8 te) 45 () - 6 = te)
Costa Rica 205 8 848 6 214 17 162 34
Cuba 31 15 159 22 100 79 41 36
Dominica 12 1 59 2 13 2 2 i)
Dominican Republic 20 ie) 125 te) = 22 = 15
El Salvador 135 1 2450 i°) 73 4 23 0
Greenland (Denmark) = te) = () = t) = t)
Grenada 14 0 50 1 12 1 3 te)
Guadeloupe 10 2 = 1 = 2 = 2
Guatemala 184 4 480 0 231 19 88 25
Haiti 20 0 = 0 = 29 = 17
Honduras 173 1 = 1 152 11 56 9
Jamaica 22 3 159 25 = 25 = 18
Martinique 9 0 53 1 = 3 = ty)
Mexico 439 136 961 88 717 368 284 169
Montserrat 8 i) 43 1 = 2 = tt)
Netherlands Antilles = 0 = i°} = 4 = i)
Nicaragua = 2 = te) 161 6 59 2
Panama 2218 11 2922 6 2226 18 164 22
Puerto Rico 13 0 94 1 46 20 22 14
St Kitts and Nevis 7 (0) 40 0 9 0 3 1
St Lucia 8 i*} 51 4 15 5 4 i*}
St Vincent and the Grenadines 9 ie) 108 2 16 3 4 0
Trinidad and Tobago 100 1 258 1 = 2 = 2
Turks and Caicos Islands = i) 184 ie) = 5 = i)
United States 346 93 650 69 = = = 122
Virgin Islands (British) = () = () = 3 - 1
Virgin Islands (US) - 0 - ie) - 4 - 1
SOUTH AMERICA
Argentina 258 47 = 21 = 63 123 37
Bolivia 280 me 1257 15 250 11 110 14
Brazil 394 68 1573 191 468 172 502 294
Chile 91 11 432 15 78 33 39 25
Colombia 359 22 1721 73 383 104 407 141
Ecuador 271 21 1435 37 337 100 343 136
French Guiana 152 1 = 1 = 1 = 2
Guyana 193 0 = 0 = 2 = 10
Paraguay 156 3 2650 0 120 4 85 4
Peru 344 46 1705 106 298 96 241 86
Suriname 187 2 - 0 = 0 = 7
Uruguay 81 () - 0) - 1 = 2
Venezuela 288 11 1308 45 - 55 = 76
OCEANIA
American Samoa 3 0 38 0 11 - 0 0
Australia 282 210 571 351 700 616 180 169
Cook Islands = te) 28 ft = = te) te)
Fiji 4 1 87 25 25 9 2 2
French Polynesia 0 () 67 25 = = 0 te)
Guam = 0 23 3 10 1 ie) 0
Kiribati = 0 15 1 = = 0 te)
Marshall Islands = 0 18 ie) 7 1 it) i?)
Micronesia, Federated States of = 3 47 18 = = 0 i)
Nauru - (o} 9 1 = = 0 0
New Caledonia if 3 116 20 32 23 0 0
New Zealand = 3 285 74 40 40 3 3
Niue 1 i) 16 0 4 ie) 0 ts)
North Marianas Islands = 0 31 3 = = 0 0
Palau = i) 48 10 22 3 1 1
Papua New Guinea 242 49 578 54 249 = 183 100
Pitcairn Islands (e) 0 19 0 = - te) tt)
Solomon Islands 47 18 163 72 57 9 15 2
Tokelau io) ie} 65 0 it (0) 0) 0
Tonga 1 i°} 39 2 6 oO i*) i)
Tuvalu = 0 9 0) = = ie) i)
Vanuatu 12 2 84 10 22 4 0 0
Wallis and Futuna Islands - te) 14 (0) = = i?) t)
Western Samoa 3 1 44 8 8 0) (0) i)
140
Higher Vertebrates
Table 13.1 Species richness and endemism: higher vertebrates (continued)
MAMMALS BIRDS REPTILES AMPHIBIANS
Species Endemic Species Endemic Species Endemic Species Endemic
known species known species known species known species
ANTARCTICA
Antarctica = 0 - 1 (0) (0) 0 0
Falkland Islands (Malvinas) = 0 63 1 te) te) t) te)
French Southern Territories = () = 1 te) to) tt) ty)
AFRICA
Algeria 92 1 192 1 = 3 = 0
Angola 276 4 872 12 = 18 = 23
Benin 188 0 630 0 = 1 = 0
Botswana 154 10} 569 0 143 2 36 1
Burkina Faso 147 1 497 0 = 3 = i)
Burundi 107 te) 633 te) = = = 2
Cameroon 297 10 848 11 = 19 = 65
Cape Verde = 0 36 4 12 10 0 0
Central African Rep 209 2 668 te) = = = 0
Chad 134 ie} 496 0 = 1 = 0
Comoros 12 2 99 9 22 3 = te)
Congo 200 1 500 0 = 1 - 1
Cote d'Ivoire 230 2 683 to) = 2 = 2
Djibouti = te) 311 (0) - - - ()
Egypt 102 4 132 0 83 1 6 0
Equatorial Guinea 184 1 392 3 = 4 = 2
Ethiopia 255 26 836 26 = 6 = 30
Gabon 190 3 617 to) = 3 = 4
Gambia 108 0 489 (0) = 1 = 0
Ghana 222 0 721 1 = 1 = 4
Guinea 190 1 529 to) = 3 = 4
Guinea-Bissau 108 (0) 376 i) = 2 ~ 1
Kenya 309 10 1067 7 187 15 88 10
Lesotho 33 ie} 288 0 = 2 = 1
Liberia 193 1 590 2 62 2 38 4
Libya 76 4 80 0 - 1 - 0
Madagascar 105 67 250 97 252 231 144 142
Malawi 195 te) 630 () 124 6 69 1
Mali 137 0 647 0 16 2 = 1
Mauritania 61 1 49 0 = 1 = 0
Mauritius = 1 102 10 = 2 2 0
Mayotte - ) - 0 15 1 = 0
Morocco 105 5 209 0 = 8 = 2
Mozambique, People’s Rep 179 2 666 i) = 5 62 2
Namibia 154 2 640 1 = 25 32 2
Niger 131 0 473 (0) = = = te)
Nigeria 274 2 831 2 >100 7 >60 1
Reunion 2 0 33 0 = 3 = 0
Rwanda 151 te) 669 (0) = 1 = 0
Saint Helena - () = 0 = pes a 0
Sao Tome and Principe 8 2 124 24 16 6 9 9
Senegal 155 1 625 0 - 1 = 1
Seychelles = 1 126 9 15 13 12 11
Sierra Leone 147 0 614 0 = 1 = 2
Somalia 171 8 639 11 193 66. 27 3
South Africa 247 27 774 7¢ 299 76 95 36
Sudan 267 7 938 0 = 6 = 2
Swaziland 47 0 381 0 106 0 39 0
Tanzania 306 12 1016 13 245 48 121 40
Togo 196 1 630 oO = 1 = 3
Tunisia 78 1 173 0 = 1 = 0
Uganda 315 4 989 3 119 2 44 0
Western Sahara 15 1 60 te) = = - 0
Zaire 415 25 1086 23 = 33 = 53
Zambia 229 3 732 () = 2 83 1
Zimbabwe 196 2 635 0 153 2 120 3
Notes: See text for general conventions adopted and sources for endemics data. Dependent islands are included with the parent territory. ? may include
marine species where data refers to mammals or reptiles, or may include non—breeding species where data refers to birds. — no data. USSR*: covers the
former Union of Soviet Socialist Republics.
141
1. Biological Diversity
Figure 13.1
Higher vertebrates: the 25 most endemic-rich countries
Hl Mammais (Birds | | Amphibians
700 |—|
sejoeds jo JOquINN
142
oF Foe
OG” Pig tO ies
Re
Country
Higher Vertebrates
Figure 13.2 Mammal richness and endemism: major countries
Ayunoy
NN a 3 RK PN 32? oh sed oe? \
en PY fot oo oon? ox? oo 290 »” oe ge yo ow” eo ed wo" evel ew we ae (ot soe
SOIWapUa-UON ae solwapuy =
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3
sejoeds jo JaquiNnN
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143
1. Biological Diversity
Figure 13.3 Higher vertebrate endemism: Asia
Hi Mammals (Birds || Amphibians
8 8
Number of species
3
&. VES PPE SE EF SS PPOs a
owe Fy es e Lop : ob 3 oe a CPS Si
Country VF we
Figure 13.4 Higher vertebrate endemism: Europe
2)
Hi Mammals {Birds | | Amphibians Reptiles
Number of species
Ho FES err POMS PEL ef SO re we “e
=\
a & Country
144
Higher Vertebrates
Figure 13.5 Higher vertebrate endemism: North and Central America
400
pd
f
8
Number of species
3
8
8
8
Number of species
i Mammals Bias Ee Amphibians
ca Mammals ee Birds ES Amphibians Es Reptiles
& PP FF SP es
we. ° oe Ct ae oh se Ago WE ra Caen
<¢
145
1. Biological Diversity
Figure 13.7 Higher vertebrate endemism: Oceania including Australia
; = Pa
L Hi Mammals §§ Birds | | Amphibians
600 | = one = ——— ~ —_ +
a
eae
Number of species
3
eC Oo S & FF PO ° © SP
AGS LIPO PEG PLP GE SE
~ & x we = we
Country
Figure 13.8 Higher vertebrate endemism: Africa and Madagascar
8
Number of species
8
a
Country ¢
LVECE SMG CT is
e
14. ISLAND SPECIES
Islands frequently have distinctive and often unique
assemblages of species. In general they have lower species
diversity than equivalent continental areas, but tend to have
elevated numbers of endemic species. The number of
species in a particular taxonomic group on a given island
and the proportion of these which are endemic appears to
depend on a wide variety of factors, both historical and
ecological. Among these are the degree of isolation, age,
size, topography and climate of the island and the biological
characteristics of the taxonomic groups concerned, in
particular their vagility (the ease with which they disperse).
Historical accident also appears to play a large part in
patterns of species occurrence on islands.
Island endemics tend to be of two types: relict species
which appear to have been more widespread in the past and
species which have evolved in isolation on the island
concerned. Relict species are generally confined to islands
which were previously part of larger land masses but which
have been isolated through processes of continental drift or
changes in sea level. Madagascar and New Caledonia are
examples of this, although, because of its size and long
period of isolation, Madagascar is perhaps more accurately
regarded as an island continent than an oceanic island. In
contrast, many island species are believed to represent the
results of adapative radiation in situ following accidental
colonisation by individuals. The biotic composition of
isolated, oceanic islands which have never been part of
larger land-masses (and are generally volcanic in origin) is
largely a result of this process. The taxa represented on
these islands are those which have (or whose ancestors had)
the capacity for long-range dispersal. Thus, at a very
general level, oceanic islands may have good representation
of, and high levels of endemism in, plants, birds and some
invertebrate groups, such as land snails and some insects,
while having low diversity of groups such as non-volant
mammals and amphibians.
Once islands have been colonised, other factors play an
important role in determining subsequent patterns of
evolution and speciation. Species which are highly vagile
tend not to speciate and diversify - this applies to, for
example, most groups of sea-birds and to strandline
vegetation. Species in these groups tend to have very wide
distributions, so that, for example, most tropical and sub-
tropical Pacific islands have essentially the same, small
number of species forming their shoreline vegetation. In
contrast, groups such as the rails (Rallidae), pigeons
(Columbiformes) and tortoises (Testudinidae) which are
essentially terrestrial but which have the capacity for long-
range dispersal will tend to form separate species on islands
or island groups which they successfully colonise. The
degree of speciation which occurs on islands subsequent to
colonization appears to be highly dependent on habitat
diversity, which is itself dependant on the size, topography
and climate of the island. Thus, low-lying oceanic islands,
such as coral atolls, tend to have low diversity and low
rates of endemism for most groups, while montane
(generally volcanic) islands tend to have much higher
species diversity and rates of endemism. As with continental
ecosystems, other factors being equal, species diversity
increases with decreasing latitude.
147
Island Species
Island - especially oceanic island - biotas tend to share
similar features, such as gigantism in plants and reptiles,
dwarfism in large mammals (although most examples of this
are extinct) and flightlessness in birds. These may arise
from the disharmonic colonisation of islands and the
subsequent evolution of plants and animals in isolation
(Bramwell, 1979). Of particular importance to conservation
are those factors which appear to lead to an increasing
extinction-pronenessamongst island species (discussed more
fully in Chapter 16). These are largely related to the
evolution of island species generally in the absence of large
terrestrial ‘predators’ - for plants these being grazing
mammals, for animals these being carnivores. This helps
explain the often catastrophic effect of the introduction of
animals such as rats, rabbits, goats, pigs and cats on native
island biotas.
This report discusses two important island groups - plants
and land snails - in some detail. Available data on these two
groups has been collated in Tables 14.1 and 14.3.
It is impractical, in a global approach, to treat each island
individually, but appropriate to consider them in groups. In
this report, we have mainly followed the classification of
islands into 147 units made by the International Working
Group on Taxonomic Databases for Plant Sciences (TDWG)
(Hollis and Brummitt, in press).
For plants, coverage of true oceanic islands is reasonably
complete. Most important continental shelf islands other
than those of the Sunda Shelf and New Guinea have also
been included.
Where complete datasets are available for particular islands,
regression analysis shows a moderately close relationship
between numbers of endemic plants and snails (see
Fig. 14.1) but no clear relationship between snails and birds
or between plants and birds (bird data not shown).
Figure 14.1 Islands: relationship between
plant and snail endemism
2 100
See.
q 50
2
<
2 20+
:
iy Me
Nr * A &
t Nae
2b Ye
WZ 1 1 4 a i He
0,01 0.03 0.14 03 4 3
Endemic plants (1000 species)
1. Biological Diversity
PLANTS ON OCEANIC ISLANDS
The number of endemic species and the proportion of the
flora that is endemic varies considerably from island to
island and appears to depend on a number of the factors
outlined above. On some island groups, like the Hawaiian
Islands, the flora can be described as consisting mainly of
‘endemics and aliens’; here the endemic species form 89%
of the native flora (Wagner et al., 1990). However, on
other islands, such as those of the Caribbean, the endemics
form only a small element in a diverse flora of
predominantly widespread continental species.
The extent to which island endemic floras consist of relict
species tends to be a matter of speculation. Greuter (1979)
suggests that about half the flora of Crete, for example, is
of the relict element. Palaeontologists have found fossils of
some Canarian endemics in southern Europe and south
Russia; these species include the famous Dragon Tree
(Dracaena draco), and the dominant species of the Canarian
laurel forests, at present a vegetation type now only found
in parts of the Canaries, Madeira and to a lesser extent the
Azores. The implication is that this remarkable type of
forest, now endangered in much of its range, once covered
much of the Mediterranean Basin in the Miocene Period, up
to 20 million years ago (Bramwell and Bramwell, 1974).
The relict species include an extraordinary array of endemic
monotypic genera and even families. Monotypic families
(i.e. families with only one species each) on islands include
Lactoridaceae (Lactoris fernandeziana) on Juan Fernandez,
Dirachmaceae (Dirachma socotrana) on Socotra, and
Degeneriaceae (Degeneria vitiensis) on Fiji. All are
threatened species and, in consequence, threatened families.
In contrast, many of the endemics have evolved in isolation
on islands. In the Canary Islands, for example, adaptive
radiation of colonists has led to over 30 endemic species in
each of the genera Echium (Vipers Bugloss), Limonium (Sea
Lavender) and Aeonium. The most outstanding example of
diversification and adaptive radiation in the plant world is
the Hawaiian Islands, where some genera, such as Cyanea
and Cyrtandra, have over 50 endemic species. Wagner et
al. (1990) propose that 469 Hawaiian species, in 20 large
genera, evolved from only 26-32 different colonists, clearly
showing the scope of the evolutionary capacity of isolated
islands. The species that result from adaptive radiation tend
to be difficult to classify, often with much hybridisation
between the various species. As with the Galapagos finches,
which helped Darwin develop the theory of evolution and
natural selection, these series of evolving and evolved island
endemics are of great importance to science.
One of the most extraordinary features of island plants is
the phenomenon of gigantism. A group of plants that is
otherwise herbaceous and often weedy is represented on
some islands as tall shrubs or trees. For example, the
endemic species of Vipers Bugloss (Echium) and the Sea
Lavenders (Limonium) in the Canary Islands include woody
shrubs with stems several metres high. Some of the most
remarkable examples are the tree daisies (Compositae) on
St Helena in the Atlantic Ocean and on the Juan Fernandez
islands off Chile.
148
In assessing the importance of islands for conservation of
the world’s plants, the best single measure is simply the
number of species endemic to the island or island group. In
virtually all cases, for plants, estimates of some kind are
available, varying from counts made from detailed floristic
analyses to estimates by knowledgeable botanists. This is
one of the few datasets on biodiversity, at least for plants,
that is complete to a reasonable standard of accuracy
worldwide.
Table 14.1 lists the islands and island groups of the world
of less than 120,000km? in size in declining order of
endemic plant species (covering flowering plants,
gymnosperms and ferns). This gives a rough guide to the
importance of each for botanical conservation. Of the
greatest importance are those three islands with over 1,000
endemic plant species each - Cuba with 3,233, New
Caledonia with 2,480 and Hispaniola (the Dominican
Republic and Haiti) with 1,800. The Hawaiian Islands were
previously included but the first comprehensive and
complete account of the plants has reduced the number of
endemics to below the thousand.
The number of endemics is an effective measure of the
importance of individual islands or island groups for plant
conservation worldwide. However, this very simple
approach is less appropriate where a significant part of the
endemic flora is shared between two or more of the island
groups used. For isolated islands or island groups like St
Helena, Juan Fernandez and the Hawaiian Islands, the
number of endemics shared with other island groups is very
small. But in the Lesser Antilles (the Leeward and
Windward Islands) in the Caribbean the shared endemics
form a considerable proportion of the endemic flora as a
whole. This is partly a consequence of the geographical
classification used; because many of the islands are
individual nation states, the classification tends to treat each
individual island as a single unit, rather than to cluster them
together, as with, for example, the Galapagos Islands or
Canary Islands. It is partly a consequence of the geography
and biology of the islands; the islands tend to be close
together, and have similar climates and land forms. Also,
because of their proximity to the Greater Antilles (Puerto
Rico, Cuba, Hispaniola and Jamaica), and to Central and
South America, there are numerous shared species both
within the Lesser Antillean chain and between various
islands of the chain and neighbouring areas.
The completion of the Flora of the Lesser Antilles (Howard,
1989) has permitted an analysis of the endemics of this
region. The results are given in Table 14.2, below, and in
the accompanying map (Figure 14.2). The table shows the
number of plant endemics with different patterns of
distribution recorded in the Flora. As can be seen, only 107
of the 327 species endemic to the Lesser Antilles as a whole
are endemic to single TDWG units (and so are included in
Table 14.2). The highest number of endemics for any island
is 25 on Guadeloupe, which is also home to a further 111
Lesser Antillean endemics.
The map shows the distributions of 190 of the 327
endemics. Most of the combinations of islands that had only
one or two endemics were omitted from the map, as were
all combinations of over five islands, as being too complex
Island Species
Table 14.1. Oceanic islands in declining order of endemic plant species
ISLAND NO. OF ENDEMIC DATE OF ISLAND NO. OF ENDEMIC DATE OF
PLANTS INFORMATION PLANTS INFORMATION
Cuba 3233 1991 American Samoa 27 1982
New Caledonia 2480 1991 Virgin Is (US) ea) 1974
Hispaniola 1800 ',* 1984 Sardinia 26 1991
Jamaica 894 1988 Guadeloupe 25 1974-89
Taiwan 892° 1982-91 Tonga 25 1991.
Hawaii 850 ? 1990 Martinique 24 1974-89
Fiji 700 ® 1984 Tuamotu Is 20 *® 1931-5
Canary Is 593)* 1990 Pitcairn Is 19° 1983
Caroline Is 293) 9% 1979, 82 St Vincent 19 1974-89
Socotra 267 1991 Netherlands Antilles 7-19 ?
Mauritius 246°” 1991 Cayman Is 18! 1984
Puerto Rico 234 1982 Annobon 17/ 1973
Trinidad-Tobago 215 1981 Christmas | 17 1980s
Ogasawara-Shoto 152 1978 Coco, Isla del 15° 1966
Vanuatu 150 1975 Bermuda 14 1991
Galapagos Is 148 1980s Guam 14 1991
Andaman Is 144 1989 Dominica 12 1974-89
Tubuai Is 140 '° 1984 Falkland Is 12 1991
Comoros 136 1917 Gambier ih! 1974
Juan Fernandez 123 1991 St Lucia 11 1974-89
Réunion 120 °, 1991 Ascension | 10 1991
Madeira 118 1980s Kazan Retto 9 1991
Bahamas 112 1982 Turks and Caicos Is 9 1982
Sao Tome 108 1944 Auckland Is 6! 1985
Marquesas Is 105 1931-35 Easter | 6 1990
Cape Verde 92 1974-79 Antigua-Barbuda Bes 1938
Cyprus 90 1977-91 Maldives 5 1961
Lord Howe | 84 1991 Malta 5 1991
Northern Marianas 81 34 1979, 82 Wallis and Futuna 51 1977
Nicobar Is ene 1989 Antipodean Is 4 1981
Balearic Is 70 1991 Grenada 4 1974-89
Seychelles 63? 1991 Selvagens 4 1980s
Western Samoa 577 ? Barbados 3 1974-89
Azores 49 1980s Campbell Is eh 1961
Bioko 49 1978 Cook Is 3 1991
St Helena 46 1991 Macquarie | g\ 1960
Corsica 45 1991 Montserrat 2 1974-89
Rodrigues 45 1991 St Kitts-Nevis 2 1974-89
Aldabra 43 * 1980 St Martin-St Barthélémy 2 1974-89
Sicily 41 1991 Marion and Prince Edward Is 1-2 1989
Tristan da Cunha 40 1965, 81 Anguilla 1 1974-89
Chatham Is 36 1991 Antigua-Barbuda 1 1974-89
Norfolk | 36 1991 Kerguelen Is ihe 1975
Principe 35 1944 Nauru 1 ?
Solomon Is 30 1991 Netherlands Leeward Is 1 1974-89
Sources: Compiled from numerous sources. See Davis, S. et al. 1986. Plants in Danger: what do we know? for many pre-1986 references.
Notes: ' Omits ferns; 7 Omits ferns and gymnosperms; * Omits monocotyledons; * Includes subspecies and varieties; * Estimated from a given
percentage of endemism; © A slight underestimate as omits endemic species treated as infraspecific level in the WCMC plants database; ’ Certainly
an underestimate; * Covers Haiti and Dominican Republic; * An underestimate as omits full treatment for families not yet covered in the Flore des
Mascareignes; © Probably an underestimate as only for Great Nicobar Island; * Omits the coralline islands, which are listed under Aldabra.
to display graphically. However, the map does show the Alfred Russel Wallace 1892. Island Life, 2nd edn.
broad pattern of plant endemism in the region, and provides Maemillan, London. 563pp.
a convincing argument for a regional approach.
Global distribution of snail diversity
LAND SNAILS
Recent estimates of world land snail species richness
"If we take the whole globe, more species of land shells are suggest a total of between 30,000 and 35,000 species
found on the islands than on the continents." (Solem, 1984).
149
1. Biological Diversity
Table 14.2 Plant endemism in the Lesser Antilles
NO OF SPECIES NO. OF TDWG UNITS NO. OF SPECIES MAPPED
107 occur in 1 unit 107
55 occur in 2 units 48
47 occur in 3 units 20
40 occur in 4 units 15
18 occur in 5 units 8
60 occur in > 5 units ie)
TOTAL ENDEMICS 327 190
Source: Howard, R.A. 1974-89. Flora of the Lesser Antilles. 6 vols. Endemics counted by Hugh Synge, 1991.
Notes: Geographical units used: (from north to south) Anguilla, St Martin-St Barthélémy, Netherlands Leewards (Saba and St Eustatius), St Kitts-
Nevis, Barbuda-Antigua, Montserrat, Guadeloupe (including Marie Galante, Les Saintes and Le Désirade), Dominica, Martinique, St Lucia, St
Vincent, Barbados, Grenada. The TDWG classification divides the Grenadine islands between St Vincent and Grenada, and so records for ‘The
Grenadines’ in the Flora have been disregarded.
Figure 14.2 Plant endemism in the
Lesser Antilles
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Species richness and endemism in land snails tend to be
closely correlated; areas with high diversity generally have
high endemism. This close relationship is shown graphically
in Fig. 14.3 (the named islands below the line have fewer
endemics than expected). On several islands with high snail
diversity all the native species are endemic and the only
non-endemics are those introduced by man. Land snail
richness and endemism are distributed very unevenly around
the world, and tend to be highest on islands and in
mountains.
A major problem in discussing mollusc richness and
endemism is the lack of information for several regions of
the world, notably Asia, the Neotropics and the Nearctic;
some continental tropical areas are particularly under-
recorded and new data could significantly change the
current picture of land snail diversity patterns.
Although islands often have highly diverse habitats, not all
islands have rich snail faunas. Work in Melanesia (Peake,
150
1969), and on the Greek islands in the Aegean Sea,
suggests that there is a direct correlation between island size
and snail species richness. Other work in the Pacific
suggests that this relationship is not always a simple one,
and Solem (1973) (also Peake, 1981) concluded that highest
diversities are found on islands about 15-40km? in area and
with an elevation of over 400m. Altitude is thus an
important factor, and atolls, for example, do not have high
snail richness or endemism.
There is some indication that isolation is also an important
factor. The island with the greatest number of species is
Rapa, one of the smallest and most remote islands in
French Polynesia. The location with the highest known snail
species richness (i.e. greatest number of species per unit
area) is Manukau Peninsula in North Island, New Zealand,
where 82 species have been found in a small area.
There is some evidence that although islands often have
remarkably high diversity and abundance (in the absence of
human impact), their snail faunas are often not ‘saturated’
and additional snail species could survive. Evidence for this
is seen from work in Madeira and on the Greek Islands,
where humans have introduced species but the numbers of
endemic species have stayed the same (Solem, 1984).
Correlation of land snail diversity with other species
Patterns of land snail diversity and endemism are generally
considered not to correlate strongly with those for other
groups of animals, particularly higher vertebrates. Available
data for islands show a marked positive correlation between
numbers of endemic plant species and endemic molluscs
(Fig. 14.1), but not between molluscs and birds. There is
a lack of data on mollusc faunas of tropical continental
areas, and it is thus difficult to make more general
statements.
Solem (1984) draws attention to the following islands as
known or believed to be important for snails:
e Reasonably well studied large snail faunas on the small
high islands of Micronesia, Melanesia, Polynesia,
Indonesia, Philippines, Mascarenes, Antilles, Madeira.
e Surveys or studies under way suggest important snail
faunas in Japan, Oahu, Tahiti, New Caledonia, New
Island Species
Table 14.3. Land snails: species richness and endemism on islands
TOTAL SPECIES ENDEMIC SPECIES % ENDEMICS
ATLANTIC
Atlantic (Macaronesian) Islands
Azores 98 41 41.8
Canary Is 181 141 77.9
Cape Verde Is 37 16 43.2
Madeira 237 171 88
Selvagens 1 1 100
Mid—Atantic Islands
Annobon (Pagalu 9 7 777
Bioko (Fernando Po) 6 c.4 c.66.6
Principe 26 15 57.7
Sao Tomé 26 19 73
St Helena c. 31 c. 25 c. 80
South Atlantic
Falkland Is 1 0 ts)
Northern European Islands
Faeroe Is 20 0 to)
Iceland 35 ie) 0
Svalbard 0 0 0
MEDITERRANEAN
Corsica c. 100 c.10 c.10
Cyclades 88 >20 c. 23
Malta c. 46 Cul, c.15
Pityuse Is 36 4 11
Sardinia = 21 =
INDIAN OCEAN
Aldabra c.9 c.4 c. 44
Adamans and Nicobars 81 75 93
Anjouan 58 = =
Comoros (inc. Mayotte) 136 = -
Grand Comore 37 = =
lle Europa 6 0-3 0-50
Mascarene Is 145 127 87.6
Mayotte 90-95 32-41 29-39
Mauritius 109 77 70.6
Moheli 18 = =
Réunion 40 16 40
Rodrigues 25 15 60
Seychelles c. 57 c. 24-26 c. 44
Socotra 49 46 94
» Sri Lanka c. 265 = c.95
Madagascar 380 361 95
CARIBBEAN
Barbuda 10 (e) (e)
Barbados 37 c.5 c.7
Cuba c. 600 = -
Guadeloupe 53 9 17
Jamaica 400-450 = 80-95
Martinique 37 15 c. 40
St Bethelemy = 0 =
St Martin c.36 0 0
Saba 14 0 0
Puerto Rico >85 = =
Mona 12 6 50
PACIFIC
Eastern
Japan 492 c. 487 99
Southwestern
Fiji 60 = =
Viti Levu 58 = =
Lakemba 22 - =
Karoni 20 = =
Mothe 13 = =
New Caledonia 300 c. 299 99
Tutuila = 8 =
Upolu 44 = =
Solomon Is 200-270 = =
Tikapia 16 7 44
Vanuatu 58 57 98
Wallis 15 te) te)
Futuna 21 ere c.5
South—Central
Henderson c.18 3 c.16
Tahiti 80 c.72 90
Rapa >105 >105 100?
North and North—Central
Hawaiian Is c. 1000 c. 1000 c. 99.9
Oahu 395 c. 387 98
Kauai 70-80 71 99
Maui 167 - =
Lanai 54 = =
Molokai 126 = =
Hawaii 128 = =
Pacific Islands off Central & South America
Galapagos c. 90 >66 c.73
Juan Fernandez Is 23 23 100
Australia and New Zealand
New Zealand c. 1000 = =
Kermadec Is c. 20 ¢c. 20 =
Lord Howe | c. 85 c. 50 c. 60
Norfolk | 84 c. 84 100
Source: table provided by Susan M. Wells (IUCN/SSC Mollusc Specialist Group)
Notes: c. approximated figure. > figure is minimum estimate.
151
1. Biological Diversity
Figure 14.3
300
100
30
T
Endemic species
Island snails: relationship between species richness and endemism
CORSICA
46 MALTA
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4K PITYUSE
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3) 10 30 100 300 1,000 3, 000
Total species
Zealand, Madagascar, Madeira.
e Poor information available but almost certainly important
islands: Hispaniola, Cuba, Jamaica, New Guinea.
Some of these areas, particularly the small high islands, do
not have particularly high diversities of vertebrates.
Ecology of snails and diversity patterns
Snails that have colonised islands and subsequently
speciated tend to be those that are good at dispersal and
thus tolerant of stress: the key factors are the presence of a
shell to resist desiccation (few slugs are found on islands),
and ovoviviparity. On most islands which have high snail
diversity, snails are largely confined to the interiors and
more mountainous regions and are often forest species
restricted to primary forest.
Viable populations of certain snail species appear to be able
to exist in very small areas over very long periods of time;
this must contribute to maintenance of high species
richness. Factors favourable to land snail speciation and the
persistence of diverse faunas are: (1) a stable and moderate
water supply providing a moist habitat (without either
torrential downpours or arid periods), (2) deep litter, (3) a
topography of gullies along streams sheltered from
prevailing winds, (4) lack of disturbance by man, (5) small-
scale vegetation changes e.g. as a result of climatic
variation, (6) little predation. Such criteria are found on
many volcanic islands and in mountains.
152
Environmental conditions that are not optimal for snails
include: (1) certain types of forest such as rain and
monsoon, which may have little litter, an overabundance of
rain, acidic soils and seasonal climates; (2) grassland (which
may however provide local conditions leading to high
abundance); (3) deserts (except where there are mountain
refugia).
Threats and extinctions
Known extinctions of island land snails are listed in Chapter
16. Solem’s work in the Pacific (Solem, 1976, 1983) gives
some idea of the rates of extinction that may be taking
place. The endodontoid snails (Families Endodontidae,
Charopidae, Punctidae) are tiny tropical snails, only a few
millimetres in diameter and are the most diverse group in
the Pacific where over 600 species have been described.
Over 100 may have become extinct this century; they are
mainly ground dwellers in primary forest and are threatened
by habitat loss and introduced ants (that prey on the eggs).
Other important island families are entirely or largely
arboreal, such as the Partulidae. This family is restricted to
the Pacific and comprises about 120 species, most of which
are probably threatened. Most is known about the Partula
of the Society Islands, where they are threatened
particularly by the introduced carnivorous snail Euglandina
rosea. Many populations of achatinelline snails in Hawaii
have been lost because of over-collecting and habitat
modification; these species are rendered highly vulnerable
to extinction because of very low lifetime fecundity (6-24)
(Hadfield, 1986). Tillier (in litt., 10 Sept. 1991) says that
from his experience (Caribbean, New Caledonia) the island
land snails most at risk are those in dry lowland forests
which may be lost to cattle grazing or development more
rapidly than upland forest.
In New Zealand at least, and probably elsewhere, the native
snails are totally dependent on native plant associations for
survival. In this country the rate of extinction is apparently
fast outstripping the rate of description of undescribed
species, many of which are ‘spot’ endemics, restricted to
tiny alpine localities or areas of limestone outcrop (Climo
et al., 1986).
References
Bramwell, D. 1979. Introduction. In: Bramwell, D. (Ed.), Plants and
Islands. Academic Press. Pp.1-10.
Bramwell, D. and Bramwell, Z. 1974. Wild Flowers of the Canary
Islands. Stanley Thornes (Publishers), London.
Climo, F.M., Roscoe, D.J. and Walker, K.J. 1986. Research on land
snails in New Zealand. WRLG Research Review No. 9. Wildlife
Research Liaison Group, Wellington, NZ. 28pp.
Greuter, W. 1979. The origin and evolution of island floras as
exemplified by the Aegean Archipelago. In: Bramwell, D. (Ed.),
Plants and Islands. Academic Press. Pp.87-106.
Hadfield, M.G. 1986. Extinction in Hawaiian achatinelline snails.
Malacologia 27(1):67-81.
153
Island Species
Hollis, S. and Brummitt, R.K. (in press). World Geographical Scheme
Sor Recording Plant Distributions. International Working Group on
Taxonomic Databases for Plant Sciences (TDWG) and Hunt
Botanical Library, Pittsburg.
Howard, R.A. 1974-89. Flora of the Lesser Antilles. 6 vols. Endemics
counted by Hugh Synge, 1991.
Peake, J.F. 1969. Patterns in the distribution of Melanesian land
Mollusca. Philosophical Transactions of the Royal Society B
255:285-306.
Peake, J.F. 1981. The land snails of islands - a dispersalist’s view
point. In: Forey, P.L (Ed.), The Evolving Biosphere. British
Museum (Natural History), Cambridge University Press. Chapter
19.
Solem, A. 1973. Island size and species diversity in Pacific island land
snails. Malacologia 14:307-400.
Solem, A. 1976. Endodontoid land snails from Pacific Islands
(Mollusca: Pulmonata: Sigmurethra). Part I. Family Endodontidae.
Field Museum of Natural History, Chicago. 508pp.
Solem, A. 1983. Endodontoid land snails from Pacific Islands
(Mollusca: Pulmonata: Sigmurethra). Part II. Families Punctidae
and Charopidae, zoogeography. Field Museum of Natural History,
Chicago. 336pp.
Solem, A. 1984. A world model of land snail diversity and abundance.
In: Solem, A. and Bruggen, A.C. van (Eds), Worldwide Snails.
E.J. Brill/W. Backhuys, Leiden. Chapter 1, pp.6-22.
Wagner, W.L. et al. 1990. Manual of the Flowering Plants of
Hawai'i. 2 vols. University of Hawaii Press, Bishop Museum
Press.
Chapter based on plant account provided by Hugh Synge
and snail account supplied by Susan M. Wells (and the
ITUCN/SSC Mollusc Specialist Group.)
1. Biological Diversity
15. CENTRES OF SPECIES DIVERSITY
INTRODUCTION
A principal goal of conservation activity is to ensure the
long-term survival of as many species as possible.
Traditionally, most resources available have been allocated
to single ‘flagship’ species, either through in situ measures
or through ex situ captive breeding efforts. Often these are
large, charismatic species which generate considerable
public interest. Habitat destruction and modification are the
most important factors now affecting species survival and
although conservation initiatives focused on single species
may protect a particular organism’s habitat, and by
extension a host of other associated species, they do not
necessarily conserve those habitats which contain the most
species.
Biodiversity is not distributed uniformly across the globe:
some habitats, particularly tropical forests among terrestrial
systems, possess a greater number or density of species
than others. Thus a 13.7km? area of the La Selva Forest
Reserve in Costa Rica contains almost 1,500 plant species,
more than the total found in the 243,500km? of Great
Britain, while Ecuador harbours more than 1,300 bird
species, or almost twice as many as the USA and Canada
combined (Myers, 1988). Given the budgetary constraints
on conservation and the competing demands of other forms
of land-use, some system is necessary for identifying the
areas in which a certain allocation of effort will maximise
species survival. It is widely accepted that the identification
and prioritisation of important centres of biodiversity are
necessary at both the national and the global scale. A
number of methods by which such areas could be
determined have been suggested.
METHODS OF DETERMINING AREAS OF
CONSERVATION PRIORITY
Overall species diversity
The simplest method of suggesting target areas for
conservation action is to identify countries with the highest
number of species (greatest species richness). For example,
Mittermeier (1988) and Mittermeier and Werner (1990)
recognised that a very small number of countries situated
mainly in the tropics possess a large fraction of the world’s
species diversity, and introduced the concept of
‘Megadiversity Countries’ which, they suggested, merit
special international attention. McNeely et al. (1990) used
country species lists of vertebrates, swallowtail butterflies,
and higher plants to identify 12 such megadiversity
countries: Mexico, Colombia, Ecuador, Peru, Brazil, Zaire,
Madagascar, China, India, Malaysia, Indonesia and
Australia. Together these countries hold up to 70% of the
world’s species diversity in these groups. This approach is
relatively simple in that it involves species inventory within
a given geopolitical boundary; it also recognises that
conservation action is managed at the country level. One
drawback to this approach, however, is that it fails to take
into account the uniqueness of the fauna and flora of a
country or region. There may be considerable overlap in
species composition between different regions with high
species numbers, particularly if they are situated close to
154
one another geographically. Taking mammal species in two
of the megadiversity countries listed above as an example,
271 species of mammal (excluding Cetacea) have been
recorded from Ecuador and 344 from neighbouring Peru,
but 208 of these are common to both countries. In addition,
high diversity regions may contain large numbers of very
widely distributed species which are currently neither
threatened nor otherwise of special conservation concern.
Endemic species diversity
An alternative approach is to identify areas with the greatest
numbers of ‘endemic’ or ‘restricted-range’ species. An
endemic species is one restricted to some given area, which
might be a mountain top, a river, a country or continent. In
this context, the assessment is often based on single-country
endemics, or on some small identifiable region within a
country. At the global level these are areas of high
conservation priority because if unique species are lost they
can never be replaced. Although not biologically
meaningful, the choice of country boundaries for assessing
endemicity is of great practical significance because
conservation action is usually administered at the national
level.
An important study that attempted to use endemic plant
species to identify areas of global conservation concern was
that of Myers (1988). Focusing on tropical forests, Myers
identified 10 regions or ‘Hot Spots’ that are characterised
by high concentrations of endemic species and are
experiencing unusually rapid rates of habitat modification or
loss (Table 15.1). These 10 areas cover only 292,000km?,
or 0.2% of the Earth’s land surface, and comprise 3.5% of
the remaining primary forest. Together, however, they
harbour 34,400 endemic plant species (27% of all tropical
forest species and 13% of all plant species worldwide).
In a subsequent publication, Myers (1990) identified a
further eight terrestrial hot spots, four in tropical forest
areas and four in Mediterranean-type areas (Table 15.1).
Together these contain 15,555 endemic plant species, or 6%
of the world’s total, in 454,400km? or 0.3% of the world’s
land area. This second selection of eight areas are therefore
not nearly as rich in endemic species as the first 10,
containing only 45% as many plant species in an area one
and a half times as large. In total these 18 sites contain
approximately 49,955 endemic plant species, or 20% of the
world’s plant species, in just 746,400knr, or 0.5% of the
Earth’s land surface.
Despite its limitations (e.g. the difficulty of quantifying
threats to the existing habitat, and the paucity of
distributional information available for many of the world’s
plant species), Myers’ work is an important step towards
determining areas where conservation requirements are
greatest and where the potential benefits from conservation
measures would be maximised.
From the wider conservation perspective, the question of
interest is whether levels of endemism in one taxon are
correlated with those in others. If endemism follows similar
patterns for different taxa, then conservation measures
Table 15.1
REGION HIGHER PLANTS
Cape Region (South Africa) 6,000?
Upland western Amazonia 5,000'
Atlantic coastal Brazil 5,000'
Madagascar 4,900'
Philippines 3,700!
Borneo (north) 3,500!
Eastern Himalaya 3,500'
SW Australia 2,830?
Western Ecuador 2,500'
Colombian Chocé 2,500!
Peninsular Malaysia 2,400'
Californian floristic province 2,140?
Western Ghats (India) 1,600?
Central Chile 1,450?
New Caledonia 1,400'
Eastern Arc Mts (Tanzania) Babe
SW Sri Lanka 500?
SW Céte d’Ivoire 200?
TOTAL 49,955
Centres of Species Diversity
Numbers of endemic species present in 18 ‘Hot Spots’
MAMMALS REPTILES AMPHIBIANS SWALLOWTAIL
BUTTERFLIES
15 43 23 fo)
= 2 c. 70 -
40 92 168 7
86 234 142 11
98 120 41 23
42 69 47 4
- 20 25 -
10 25 22 fe)
- - 2
137 111 {o}
25 7 fe)
15 15 16 {o}
7 91 84 5
2 21 {o} 2
20 - 49 3
4 - - 2
3 - 2 {0}
375 892 737 59
Sources: For plants, Myers (1988', 19907); for animals, miscellaneous sources (WCMC).
Notes: - indicates no data yet available. All regions are classed floristically as tropical forest, with the exceptions
of four regions which have Mediterranean-type floras, i.e. Cape Region South Africa, SW Australia, Californian floristic province and Central Chile.
focused in areas of high endemism will generate enhanced
returns in terms of overall biodiversity conservation.
Myers’ botanical hot spots are undoubtedly good sites to
conserve endemic plants, and they often contain high
numbers of endemics among other groups. There are
exceptions, however, and the strength of such relationships
remains to be investigated. Area, size, scale, and the
biogeography of different taxa will be among the important
variables.
Bibby ef al. (1992) examined available data for other
groups to compare with bird data, and showed that
endemism at least among larger vertebrates is often, though
not always, related. Countries with high numbers of
endemics in one vertebrate group often also have high
numbers of endemics among other vertebrates (see Table
15.2). Statistically, numbers of mammals and birds, and of
mammals and reptiles, correlate quite closely. Country size
is probably an important factor underlying these
correlations: larger countries tend to have larger numbers
of species and also larger numbers of endemic species of
each taxon.
Even if associations do exist between levels of endemicity
in different taxa, care must be exercised in their
interpretation and application since correlations are merely
generalisations. For example, Table 15.1 shows that while
there may be some broad similarities amongst endemic
species numbers in different vertebrate and plant taxa, there
are significant discrepancies. Thus, although the Colombian
Choc6é has high numbers of endemic reptiles and
amphibians (137 and 111 respectively) it has relatively few
155
endemic mammals (8); and the Cape Region of South
Africa, which has the highest number of endemic plant
species (6,300) has only 15 endemic mammals. Overall
conservation priorities should therefore be based on a
synthesis of detailed analyses of different taxonomic groups,
not an analysis of the pattern of endemicity in just one
taxon.
Critical faunas analysis
Whether simple species richness or levels of endemism are
initially used to assess the biological importance of sites,
the concept of ‘complementarity’ and its application in
‘critical faunas analysis’, first introduced by Ackery and
Vane-Wright (1984), is increasingly used to determine
conservation priorities objectively. In this approach the
entire set of taxa within the group under consideration, e.g.
single-country endemic amphibians, constitutes the
‘complement’. The single most important site for
conservation is that at which the greatest proportion of the
complement is represented. The portion of the complement
not included is called the ‘residual complement’. The
priority for second site selection can be determined by
identifying the site that adds the greatest proportion of the
residual complement to the initial choice. The process can
be continued in a step-wise sequence until all sites have
been considered and allocated a priority. The advantage of
this process is that it produces an objective and optimised
selection sequence, against which the performance of any
other (sub-optimal) sequence can be judged for its relative
efficiency in representing total biodiversity.
1. Biological Diversity
Table 15.2 Countries rich in endemic land vertebrates
COUNTRY ENDEMIC TAXON
RANK
ORDER MAMMALS BIRDS REPTILES AMPHIBIANS
1 Australia 210 Indonesia 356 Australia 605 Brazil 293
2 Indonesia 165 Australia 349 Mexico 368 Mexico 169
3 Mexico 136 Brazil 176 Madagascar 231 Australia 160
4 USA 93 Philippines 172 Brazil 178 Madagascar 142
5 Philippines 90 Peru 106 India 156 Ecuador 136
6 Brazil 70 Madagascar 97 Indonesia 150 Colombia 130
az Madagascar 67 Mexico 88 Philippines 131 India 110
8 China 62 New Zealand 74 Colombia 106 Indonesia 100
9 USSR 55 Solomon Islands 72 Ecuador 100 Peru 87
10 PNG 49 India 69 Peru 95 Venezuela 76
11 Argentina 47 Colombia 58 Cuba 79 Cameroon 65
12 Peru 46 Venezuela 45 South Africa 76 Zaire 53
Source: WCMC database.
Collins and Morris (1985) performed a critical faunas
analysis at the country level, examining endemicity in
swallowtail butterflies. They found that if the five countries
with the highest numbers of endemic swallowtail species
enacted conservation plans to protect swallowtails, then
54% of the world’s total number of swallowtail species
would be conserved. If the next five countries were
included, the total protected would rise to 68%. Increments
decreased as further blocks of five countries were added,
with 15, 20, 25, 30, 35, 40 and 45 countries respectively
including 77, 90, 93, 95, 96, 97 and 99% of the world’s
swallowtails.
This type of analysis can be used to direct international and
national attention to faunistically important countries, states
or provinces. Local knowledge must however remain the
basis for more detailed conservation planning, in order to
identify precise centres of species richness and importance
within a country, and to plan a system of protection around
those centres.
Whilst earlier studies were based on species numbers alone,
more sophisticated studies of this kind are now being
developed which attempt to take into account species
turnover between sites, not only in a simple numerical sense
but by use of some taxic diversity index. Taxonomic
dispersion is the most complex but perhaps intuitively most
attractive of these, in that, given a hypothesis of the
evolutionary relationships among members of a group, it
attempts to select an even spread of taxa across the
hierarchy (see Chapter 2).
Table 15.3 shows:one application of this procedure, to
determine the priority sequence of African protected areas
for the conservation of antelopes. Serengeti National Park
(Tanzania) is the richest single site, holding breeding
populations of 24% of all African antelope species. The
highest incremental change occurs with the addition of
Kafue National Park (Zambia): together the two parks hold
38%. The addition of two further reserves, Haut Dodo
Faunal Reserve (Céte d’Ivoire) and Ouadi Rimé-Ouadi
156
Achim Faunal Reserve (Chad) brings the representation of
African antelope species diversity to over 56% in just four
protected areas.
In critical faunas analysis, if all the species in the world in
the taxon under consideration are to be conserved, and if all
species are treated as taxonomically equal, then a priori
endemics are accorded a high value in the prioritisation
sequence. Thus Ackery and Vane-Wright (1984) found that
in order to conserve all 158 species of milkweed butterflies
(Lepidoptera: Danainae) a total of 31 sites or ‘critical
faunas’ needed protection. Site selection was made starting
with the site that contained the highest number of endemics
- in this case Sulawesi. Of these 31 sites, 24 were sufficient
to protect all the narrow endemics, and a further seven
were sufficient to complete the list. In practice, even if the
conservation of 100% of the Earth’s biodiversity is the
goal, some species will of necessity be neglected. The
critical faunas approach may not always offer a sufficiently
flexible strategy for planning conservation at the global
level (Vane-Wright ef al., 1991).
Conclusion
Although species are normally used as the basis for critical
faunas evaluation or distributional analysis, other taxonomic
groupings such as genus or family can be used instead.
Different forms of weighting system can also be introduced,
so that for instance a species in a monotypic genus, such as
the Giant Panda Ailuropoda melanoleuca might be allotted
a higher conservation priority than a species with many
congeners. New measures of biodiversity are now being
developed which can take into account the genetic
distinctiveness of species based on the relative position of
species and other taxa in the classification hierarchy. For
example, Vane-Wright ef al. (1991) suggest using a ‘taxic
diversity measure’ based on the information content of
cladistic hypotheses (indicating the branching pattern of
evolution), which would provide a measure of taxonomic
distinctiveness. Bibby et al. (1992) apply a simple method
of assigning taxonomic uniqueness to endemic species based
Table 15.3
STEP DIVERSITY DIVERSITY
NO. INCREMENT % CUMULATIVE %
1 23.95 23.95
2 13.70 37.65
3 9.99 47.64
4 9.32 56.96
5 4.85 61.81
6 4.71 66.52
7 5.27 71.79
8 3.50 75.29
9 2.81 78.10
10 2.82 80.91
Centres of Species Diversity
Biodiversity scores for Afrotropical antelopes
CONSERVATION COUNTRY
AREA NAME
Serengeti NP Tanzania
Kafue NP Zambia
Haut Dodo FR Cote d’lvoire
O. Rime-O. Achim FR Chad
Yangudi Rassa NP Ethiopia
Odzala NP Congo
W. Pretorius GR S Africa
Manovo-G-St Floris NP C African Rep
De Hoop NR S Africa
Gorongosa NP Mozambique
Note: Part of the optimised priority area sequence of protected areas in terms of their potential for conservation of African antelopes, based on the
taxonomic dispersion measure and complementarity. Serengeti National Park (Tanzania) is the richest single site, holding breeding populations of
species accounting for 24% of African antelope diversity. The highest incremental addition occurs in Kafue National Park (Zambia); in combination
the two total 38%. The addition of two further reserves (one in Céte d’Ivoire; one in Chad) brings the representation of African antelope taxonomic
diversity to over 56%. (Based on data from East (1988, 1989, 1990) and Gentry (in press) and analysis of Williams (unpublished report).)
on the diversity of the genus and family to which the
species belongs.
The kinds of technique outlined above are useful tools
which enable conservation biologists to prioritise sites and
allocate scarce resources. Care must be taken to base
overall global conservation priorities on a number of taxa,
which ideally should be well-represented throughout the
world. It should, however, be remembered that the
identification of areas of high diversity is but the first step
in determining effective conservation plans. The size and
heterogeneity of the sites under consideration also have
serious implications for conservation biology through their
effects on minimum viable population sizes, stochastic
ecological effects, etc. Planners must seek to conserve
multiple populations whenever possible, to allow for chance
local extinctions. Progress in designing the protection of a
functional ecological system or set of systems has recently
been made in Australia (e.g. Margules, 1989), where
wildlife services are developing step-wise analyses intended
to take these kinds of factors into account.
Two major projects have developed and refined approaches
to the systematic identification of centres of species
endemism or diversity at the global level. The IUCN Plant
Conservation office is identifying centres of plant diversity,
and the International Countil for Bird Preservation (ICBP)
has identified centres of endemism among restricted-range
birds. The approaches and major findings of these two
projects are detailed below. Simple visual comparison of the
two world maps relating to these projects (Figs 15.1 and
15.2) shows much broad correspondence between the sites,
although there are differences in detail (e.g. more
botanically diverse areas identified in Mediterranean
fegions). The sites concerned are noted in Tables 15.6
(plants) and 15.7 (birds).
CENTRES OF PLANT DIVERSITY
The IUCN Plant Conservation Programme is at present
Carrying out a project to identify the several hundred major
Centres of Plant Diversity (CPD). These are defined as
places particularly rich in plant life which would if
protected safeguard the majority of wild plants in the world.
The book IUCN is preparing with the help of collaborators
157
worldwide will provide detailed data sheets on some 250
selected areas. It will also document the many benefits,
economic and scientific, that conservation of these areas
would bring and will outline the potential value of each for
sustainable development.
IUCN has defined the CPD ‘sites’ as of three types:
e botanically rich sites that can be defined geographically
(e.g. Mt Kinabalu in Borneo)
geographically defined regions with high species
diversity and/or endemism (such as the Atlas Mountains,
or the Cordillera Bética in Spain)
vegetation types and floristic provinces that are
exceptionally rich in plant species (such as the Amazon
rain forests and the South-West Botanical Province of
Western Australia).
The formal criteria for inclusion of sites in the Centres of
Plant Diversity project specify that each must have one or
both of the following two characteristics:
® the area is evidently species-rich, even though the
number of species present may not be accurately known
the area is known to contain a large number of species
endemic te it.
The following characteristics are also considered in the
selection: a) the site contains an important gene pool of
plants of value to man or plants that are potentially useful;
b) the site contains a diverse range of habitat types; c) the
site contains a significant proportion of species adapted to
special edaphic conditions; d) the site is threatened or under
imminent threat of large-scale devastation.
The selection is therefore based on botanical importance
rather than on degree of threat. A site that could be
considered safe one year could be severely endangered the
next. This is particularly likely in the tropics where
pressures on land continue to increase.
The site selection process involved extensive consultations
with experts in all major regions. In Africa, China, India,
North and South America, this has resulted in Workshops
at which data on lists of proposed CPD sites have been
reviewed, and the final site selection made. For the Central
Asian region, the final selection has not yet been made.
1. Biological Diversity
Figure 15.1 Centres of plant diversity: the world
e)
S)
e)
©
)
e)
fo)
+
e)
‘)
=)
N
.)
Centres of Species Diversity
Figure 15.2 Endemic bird areas: the world
js)
[e)
ie)
Ke)
ie)
oO
ie)
+
je)
ie)
oO
N
(eo)
159
1. Biological Diversity
The difficulty in selecting sites varies greatly from one part
of the world to another. In some regions the selection is
easy. In West Africa, for example, it has long been known
that the famous Tai Forest National Park is the only large
portion of rain forest in Céte d’Ivoire still intact; with over
150 plants endemic to the park, the Tai is an obvious
candidate for inclusion. Often, especially in Africa, the
Centres of Plant Diversity are mountains, like Mt Nimba
where the borders of Guinea, Liberia and Ivory Coast meet,
Mt Mulanje in Malawi, and the Air Mountains in the
Sahara. Such mountains have a wide range of diverse plant
communities but are often delimited by low-diversity habitat
making identification of sites relatively easy.
In other areas the selection is much more difficult. The
islands of Borneo and New Guinea, for example, contain
the largest floras in Asia. Virtually all the habitats are rich
in plants, but floral diversity varies from place to place in
very complex ways. As a result, it is very hard to specify
which parts of Kalimantan, if protected, would include the
most plant species. In Irian Jaya, botanical knowledge is not
yet sufficient to say with any degree of confidence which
areas are richest in plant species.
In some regions, the selection of the sites that need to be
protected cannot be made on botanical criteria alone. For
example, the Atlantic forests of Brazil are reduced to 2-5%
of their original extent, and they have a quite different
complement of species to the much larger Amazonian
forests. To save their flora, as many as possible of the
surviving remnants should be protected. Where such
remnants provide two similar sites, with similar
complements of species, socio-economic considerations
rather than botanical ones will influence the decision as to
which sites might be protected. In such cases the CPD
project will identify the whole vegetation type - in this case
the Atlantic forests of Brazil - and not recommend detailed
protection strategies for the various sites within that region.
In addition to data sheets on the selected sites, the CPD
publication will contain Regional Overviews which will
describe the general patterns of vegetation and plant
distribution. Opinions will naturally vary as to the exact
choice of sites for coverage at international level, and in
order to avoid implications that only the 250 or so sites
outlined should be protected, the Regional Overviews will
also contain lists of other sites for botanical conservation,
many of a lesser priority but important nonetheless.
The ‘Centres’ concept is particularly appropriate for plant
conservation because it focuses on the plant-rich tropics. As
the map of sites (Fig. 15.1) shows, most of the 241 sites
selected so far are in the tropics, where it is not usually
possible to identify threatened plant species individually.
Botanists can, however, say which areas are rich in plants
and which are not without knowing the status of every
single species. Thus, while identifying threatened species
provides a practical approach to planning plant conservation
in most temperate countries, and on most islands,
identifying Centres of Plant Diversity is the best approach
in most of the tropics.
It is as yet unknown to what extent the sites identified as
Centres of Plant Diversity can also be described as centres
of diversity for animals; it is intended to investigate this
during later stages of the project.
All the 241 Centres of Plant Diversity selected so far are
listed in Table 15.6. As the data on degree of protection
show, many are already protected areas, such as Bwindi
(Impenetrable) Forest (Uganda), the wet tropics of
Queensland (Australia) and the Sinharaja Forest (Sri
Lanka). In virtually all cases, however, more conservation
work is needed to ensure the full complement of plants
survives intact. Below we show the areas selected for
Africa and Peninsular Malaysia, showing the application of
the approach in regions of very different size.
Centres of Plant Diversity: Africa
White (1983) recognises 17 major phytogeographic
divisions (phytochoria) for mainland Africa. Of these,
seven are classed as Regional Centres of Endemism, each
having more than 50% of its species confined to it and a
total of more than 1,000 species endemic to it. Two more
phytochoria (Afromontane and Afroalpine) are patchily
distributed on mountains. The remainder are termed
transition zones, having low species endemism and, in some
cases, very impoverished floras. These main divisions,
which cover vast areas, were used as the starting point for
selecting sites for the CPD project. In general, floristically-
rich phytochoria have been allocated more Data Sheet sites
than those with impoverished floras; however, some regions
with very high endemism, such as the Cape, will be treated
as one ‘super-site’.
Salient features of White’s phytochoria are outlined below,
and indicated in Fig. 15.3. The CPD sites are shown in the
same map, superimposed on these regional divisions and
details presented in Table 15.4. For comparison, areas of
bird endemism identified by ICBP are mapped in Fig. 15.4
and detailed in Table 15.5.
Guineo-Congolian (A)
8,000-12,000 vascular plant species; endemism very high,
80%. The tropical rain forest in west and central Africa.
Western block (Guinea) floristically distinct from central
block, mostly cleared or threatened. Gulf of Guinea
islands, especially Sado Tome, also have high endemism.
Central block (Congo) has two main centres of plant
diversity: west (especially Gabon - the most species-rich
rain forest in Africa, and Cameroon), and east (especially
Zaire). In Cameroon, forests nearer coast richer (e.g.
Korup), extending into south-east Nigeria (e.g. Oban). In
Zaire, forests near coast (e.g. Mayombe) reported to be
floristically distinct, threatened; forests on east side (e.g.
Maiko, Kahuzi-Biega, Ituri, probably Itombwe) appear to
be richer than those in centre.
Zambezian (B)
8,500 vascular plant species; high endemism, 54%.
Miombo, mopane and chipya woodland. Most diverse area
is Haut Shaba, Zaire (including Kundelungu). Zambia:
richest miombo is in wetter area near Zaire border
(extension of Haut Shaba). Angola: Huila Plateau rich in
endemics, Itigi thicket near Tanzania/Zambia border also
rich. Local endemics in Zaire on metalliferous soils and on
serpentine in Zimbabwe (e.g. Great Dyke) need protection.
Sudanian (C)
2,750 vascular plant species, most widely distributed;
regional endemism low, 35%. Woodland (mainly
Isoberlinia, Khaya).
Somalia-Masai (D)
2,500 vascular plant species; 50% regional endemism.
Acacia, Commiphora woodland. Rather homogenous;
Somalia the richest country.
Cape (E)
Fynbos, with 8,600 vascular plant species; high endemism,
60-68%. Extraordinarily rich in species and endemics.
Many important areas. Invasive species a major problem.
Karoo-Namib (F)
6,000 vascular plant species; 35-40% regional endemism.
Dwarf succulent shrubland. Unparalleled diversity of
succulents. Important centres in north (Gariep centre,
including the Richtersveld) and south (southern Namibia and
western Cape Province, South Africa).
Mediterranean (G)
4,000 vascular plant species; endemism low, 20%.
Evergreen oak forest, macchia, maquis. High Atlas the
most outstanding area botanically, many species and
endemics.
Afromontane (H)
4,000 vascular plant species; endemism very high, 75%.
Forests, afroalpine vegetation. Afroalpine vegetation
(above forest limit) especially rich in local endemics.
Eastern Arc mountains in Tanzania and south-east Kenya
have many species absent from central Africa, especially in
submontane forest. Richest mountains in central Africa
uncertain, but possibly East Kivu (e.g. Itombwe) and
Bwindi. All African mountain forests especially important
for watershed protection.
Indian Ocean coastal (M & O)
Comprising Zanzibar-Inhambane regional mosaic in north,
and Tongaland-Pondoland in south. Both with 3,000
vascular plant species and low endemism, 15-20%. Most
important are the coastal forest remnants, floristically
similar to the Guineo-Congolian, but with c. 40% of species
endemic to coastal belt, many with very restricted
distributions. In Kenya, c. 50 forest patches, most very
small. In Tanzania, number of sites uncertain, need more
fieldwork to determine which areas are key; includes Rondo
Plateau.
CENTRES OF AVIAN ENDEMISM
The International Council for Bird Preservation, in its
Biodiversity Project, has undertaken a major data collation
and analysis project to identify areas supporting
aggregations of restricted range endemic birds. This project
has served two functions: first, it applies rigorous scientific
161
Centres of Species Diversity
criteria for identifying areas of high conservation value for
birds; and second, it reviews the information on patterns of
endemism in other taxonomic groups so that the value of
birds as biodiversity indicators can be assessed. The major
results of the project are now published in Bibby ef al.
(1992).
Locality records were gathered for species with breeding
ranges below 50,000km? (about the size of Sri Lanka, Costa
Rica or Denmark). Remarkably, there are 2,608 species or
27% of the world’s birds with such small ranges. In all,
some 55,000 separate locality records of birds were
accurately geo-referenced and mapped with the aid of a
Geographic Information System.
Species of restricted range tend to occur together, for
instance on islands or in isolated areas of a particular
habitat, such as tropical montane forest. Boundaries of these
natural groupings of species have been identified
(designated as Endemic Bird Areas or EBAs). They number
221 and embrace 2,480 species, which is the vast majority
of all restricted range birds. Both the numbers of species
involved and the number of EBAs divide roughly equally
between continental areas and islands.
The tropics, with 76% of all Endemic Bird Areas, are the
most important zone and there are very few at north
temperate latitudes (Fig. 15.2). Indonesia is by far the most
important country, with 411 restricted range species of
which 339 are confined to the country. Peru, Brazil,
Colombia, Papua New Guinea, Ecuador, Venezuela, the
Philippines, Mexico and the Solomon Islands all have more
than 100.
Table 15.7 shows the political affiliation, altitudinal range
and habitats, and richness in restricted range birds of each
EBA. The size of EBAs varies considerably, from the
Northwestern Hawaiian Islands (Skm?) to the Guianas
(170,000km?). However, over 30% of EBAs have areas of
less than 10,000km? and are therefore considerably smaller
than the maximum range size allowed for any one species.
Island EBAs are generally smaller than continental EBAs.
For instance, 29% of island EBAs are smaller than
1,000km?, whereas no continental EBAs are this small. The
extent of EBAs in Africa, Middle East and Europe are
shown in Fig. 15.4 and sites are detailed in Table 15.5.
The number of restricted range bird species contained
within EBAs also varies, from the minimum of two used to
define an EBA to 67 in the Solomon Islands EBA. A large
number, 757 (29%), of these birds are threatened, and they
constitute 77% of all threatened birds. Most EBAs (85%)
have one or more threatened restricted range bird species
(see Table 15.5 for Africa and adjacent areas). The
principal habitat used by birds in the EBAs is forest (69%
of restricted range species) with smaller numbers using
scrub (12%). Other habitats such as grasslands are poorly
represented, largely because species in these habitats are
generally more widespread.
1. Biological Diversity
Figure 15.3 Centres of plant diversity: Africa
al Centres of Plant Diversity
|
0 800 1600 2400km
{<a — et ol
Notes: Data Sheet sites are shown superimposed on the main phytochoria (after White, 1983). Letter codes denote the following: (A) Guineo-
Congolian regional centre of endemism. (B) Zambezian regional centre of endemism. (C) Sudanian regional centre of endemism. (D) Somalia-Masai
regional centre of endemism. (E) Cape regional centre of endemism. (F) Karoo-Namib regional centre of endemism. (G) Mediterranean centre of
endemism. (H) Afromontane archipelago-like regional centre of endemism. (J) Guinea-Congolia/Zambezia regional transition zone. (K) Guinea-
Congolia/Sudania regional transition zone. (L) Lake Victoria regional mosaic. (M) Zanzibar-Inhambane regional mosaic. (P) Sahel regional transition
zone. (O) Tongaland-Pondoland regional mosaic. (Q) Sahara regional transition zone. (R) Mediterranean/Sahara regional transition zone.
162
Centres of Species Diversity
Figure 15.4 Endemic bird areas: Africa, Middle East, Europe
SSS
[Le]
(Source: ICBP)
Robinson projection
163
1. Biological Diversity
Table 15.4 Centres of Plant Diversity: sites in continental Africa
SITE SITE NAME COUNTRY NO. OF PLANT
NO. SPECIES
1 Sapo Forest Liberia
2 Tai Forest Ivory Coast
3 Mt Nimba Guinea, Ivory Coast, Liberia >2,000
4 Salonga National Park Zaire
5 Mayombe-Cabinda Congo, Cabinda, Zaire
6 Korup-Oban Cameroon, Nigeria 3,500
7 Dja Cameroon 2,000
8 Crystal Mountains Gabon >3,000
9 Massif du Chaillu Gabon >3,000
10 Massif de Doudou Gabon >1,000
11 Maiko Zaire
12 Bwindi (Impenetrable) Forest Uganda 1,000 taxa
13 Kundelungu/Upembe Zaire
14 Huila Plateau Angola
15 Zambesi Source Area Zambia
16 Okavango-Kwando Angola, Namibia, Botswana
17 Mbali-Mahali Hills Tanzania
18 Cape Floristic Province South Africa 8,600
19 Mt Kenya Kenya 800
20 Eastern Arc Mts: Usambaras Tanzania 1,921 taxa*
21 High Drakensberg South Africa
22 Mt Mulanje Malawi >8s00
23 Bale Mts Ethiopia >1,000
24 Mt Cameroon Cameroon 1,200
25a Karoo-Namib region South Africa 5,000 taxa
25b Gariep Centre South Africa, Namibia
25c Brandberg-Kaokoveld Angola, Namibia
26 Cal Madow Somalia >1,000
27 Hobyo Somalia <1,000
28 Limestone bush/woodland, Ogaden Ethiopia, Kenya, Somalia
29 Garamba Zaire
30 High Atlas mts Morocco
31 Pondoland Plateau South Africa
32 Rondo Plateau Tanzania
Note: Figures refer to the number of vascular plant species (if known) estimated to occur in the area. * denotes number of vascular plants so far
recorded.
Importance for other taxonomic groups
The review of other taxonomic groups suggests that the
EBAs are also of great importance for mammals, reptiles,
amphibians, molluscs, insects and plants. However, there
are gaps in data on these other groups, and additional data
on these could significantly change the conservation
evaluation of some EBAs. It seems likely, also, that in
other groups different scales of endemism may occur (more
fine-grained for various invertebrate groups and some plant
taxa, for instance), and it is worth noting that entirely
different approaches are needed to deal with non-terrestrial
endemism.
Evaluation for importance and threats
EBAs were evaluated on biological importance and threat.
The biological importance index reflects richness in
restricted range species per unit area, modified to allow for
taxonomic uniqueness of the species involved. On this
basis, ICBP assigns EBAs to three categories. EBAs which
are also significant centres of endemism for at least two
164
other taxonomic groups are upgraded by one category. This
has the overall effect of increasing the priority of EBAs
which are important for other groups: uniform quantitative
data on other groups are urgently needed to refine these
priorities. Threats to EBAs are evaluated on the proportions
of restricted-range species threatened, and the extent of
coverage by the protected areas system (see Table 15.5).
Conclusions
Bibby er al. (1992) conclude that 20% of all bird species
are confined to just 2% of the world’s land surface. The
total area of all 221 EBAs accounts for 4.5% of the land
surface. Since more widely ranging species also occur in
these EBAs, the proportion of the world’s birds that could
be conserved if EBAs were secured would greatly exceed
the 27% whose ranges are highly restricted.
Further work and follow-up
ICBP aims to promote the conservation of all 221 EBAs, in
collaboration with other international and national
Centres of Species Diversity
Table 15.5 Endemic bird areas of Africa, the Middle East and Europe
AREA NAME SIZE SPP. CONFINED SPP. OCCURRING SPP.R PA
(km?) (%)
T N Tot. T N Tot.
Sites in continental Africa
Upper Guinea forests 113,000 4 1 5 5 1 6 5.5 v/
Cameroon mountains 7,300 8 4 26 9 4 28 27.0 7)
Cameroon and Gabon lowlands 40,000 4 - 5 5 - 6 35) 22
Angola 14,000 5 4 14 6 4 15 14.5 4
North-east Somalia 41,000 2 - 5 2 - 5 5.0 (0)
Central Ethiopian highlands 37,000 2 1 4 2 1 4 4.0 {e)
South Ethiopian highlands 15,000 4 - 4 4 - 4 4.0 {e)
Central Somalian coast 1,200 1 1 2 1 1 2 2.0 (0)
East Zairean lowlands 49,000 4 1 5 4 1 5 5.0 3
Albertine Rift Mountains 44,000 8 4 37 10 4 40 38.5 12
Kenyan mountains 46,000 2 6 2 - w/ 6.5 6
Serengeti 47,000 1 3 - 1 3 3.0 43
Kenyan and Tanzanian coastal 8,800 5 - 7 6 8 7.5 7/
forests
Eastern Arc Mountains 39,000 11 2 26 13 2 30 28.0 11
South Zambia 47,000 1 1 2 1 1 2 2.0 14
East Zimbabwean mountains 4,900 - 2 2 1 2 4 3.0 7
South-east African coast 43,000 - 2 2 - 3 3 2.5 3
South-east African grasslands 60,000 2 - 2 2 - 2 2.0 (e)
Cape region 24,000 - 1 3 - 1 4 3.5 50
Sites outside continental Africa
Canary Islands and Madeira 8,100 6 - 9 6 10 8.5 30
Cape Verde Islands 4,000 1 - 4 1 - 4 4.0 ¢)
Principe 140 - 6 3 - 12 8.8 {e)
Sao Tome 860 6 1 15 9 1 21 16.8 (0)
Tristan da Cunha Islands 200 5 - 6 5 - 6 6.0 te)
Caucasus 64,000 1 2 1 2 2.0 9
Cyprus 9,300 - 2 - - 2 2.0 {¢)
lraq marshes 40,000 - - 2 - - 2 2.0 (0)
Arabian mountains 59,000 1 - 7 1 - 7 7.0 1
Socotra 3,500 1 - 6 1 - 6 6.0 [0]
Granite Seychelles 240 6 - 10 6 - 10 10.0 3
Aldabra 160 1 1 2 1 1 4 PAT 4
Comoro Islands 1,900 4 - 10 4 - 13 uve (0)
Mayotte 360 1 - 3 1 - 6 4.2 10)
West Madagascan dry forest 30,000 1 - 3 2 - 4 3.5 3
East Madagascan humid forests 112,000 13 1 17 14 1 18 17.5 5
Central Madagascan lakes 2,000 2 = 2 2 - 2 2.0 0
West Madagascan coastal 5,000 2 = 2 2 = 2 2.0 11
wetlands
South Madagascan Didiera scrub 30,000 2 j 8 2 1 8 8.0 3
Reunion 2,500 1 - 3 1 - 7 5.0 2
Mauritius 1,900 6 - 6 6 - 10 8.0 2
Rodrigues 100 2 = 2 2 - 2 2.0 te)
Key: T=threatened; N=near threatened; SPP. R=species richness; PA=coverage by protected areas.
organisations. The list of priority areas should enable a
wide range of organisations to develop both regional and
local programmes to help implement measures to prevent
mass species extinctions. These measures will range from
establishment and management of protected areas to the
sustainable use of natural resources in the centres of
endemism, and will require political and economic
collaboration at all levels. There is a need to strengthen
local data on birds and other taxa, and within EBAs, to
study habitats which are vital for the survival of restricted
165
range species. Key sites must be identified within EBAs for
the targeting of conservation resources.
References
Ackery, P.R. and Vane-Wright, R.I. 1984. Milkweed Butterflies.
British Museum (Natural History), London.
Collins, N.M. and Morris, M.G. 1985. Threatened Swallowtail
Butterflies of the World. The IUCN Red Data Book. TUCN,
Cambridge, UK and Gland, Switzerland. vii+401pp. + 8pls.
1. Biological Diversity
Bibby, C.J., Crosby, M.J., Heath, M.F., Johnson, T.H., Long, A.J.,
Stattersfield, A.J. and Thirgood, S.J. 1992. Putting Biodiversity
on the Map: global priorities for conservation. ICBP,
Cambridge, UK.
East, R. (Ed.) 1988. Antelopes. Global survey and regional actions
plans. Part 1. East and northeast Africa. TUCN, Gland.
East, R. (Ed.) 1989. Antelopes. Global survey and regional actions
plans. Part 2. South and south-central Africa. TUCN, Gland.
East, R. (Ed.) 1990. Antelopes. Global survey and regional actions
plans. Part 3. West and central Africa. UCN, Gland.
Gentry, A. (in press). The subfamilies and tribes of the family
Bovidae.
Margules, C.R. 1989. Introduction to some Australian developments
in conservation evaluation. Biological Conservation 50:1-11.
McNeely, J.A., Miller, K.R., Reid, W.V., Mittermeier, R.A. and
Werner, T.B. 1990. Conserving the World’s Biological
Diversity. UCN, Gland, Switzerland.
Mittermeier, R.A. 1988. Primate diversity and the tropical forest: case
studies from Brazil and Madagascar and the importance of the
megadiversity countries. In: Wilson, E.O. and Peter, F.M.
(Eds), Biodiversity. National Academic Press, Washington, DC.
Pp.145-154.
166
Mittermeier, R.A. and Wemer, T.B. 1990. Wealth of plants and
animals unites ‘megadiversity’ countries. Tropicus:4(1):1,4-5.
Myers, N. 1988. Threatened biotas: ‘hot spots’ in tropical forests. The
Environmentalist 8(3):187-208.
Myers, N. 1990. The biodiversity challenge: expanded hot-spots
analysis. The Environmentalist 10:243-256.
Vane-Wright, R.I., Humphries, C.J. and Williams, P.H. 1991. What
to protect? - systematics and the agonies of choice. Biological
Conservation 55:235-254.
White, F. 1983. The Vegetation Map of Africa. A descriptive memoir
to accompany the Unesco/AETFAT/UNSO Vegetation Map of
Africa.
Williams, P.H. (unpublished). Afrotropical antelopes - priority areas
for biodiversity. Progress report to The Natural History
Museum, London, WCMC and IUCN-SSC.
Text, table and maps on plant diversity supplied by IUCN
Centres of Plant Diversity Project. Text, table and maps on
bird diversity provided by ICBP Biodiversity Project.
Additional material from R.I. Vane-Wright, Biodiversity
Programme, The Natural History Museum (London).
Centres of Species Diversi
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186
Table 15.7 Endemic bird areas of the world
NAME
POLITICAL UNIT(S) ALTITUDE (m)
AFRICA, THE MIDDLE EAST AND EUROPE
Canary Islands and Madeira
Cape Verde Islands
Upper Guinea forests
Cameroon mountains
Cameroon and Gabon lowlands
Principe
Sao Tome
Angola
Tristan da Cunha Islands
Caucasus
Cyprus
Iraq marshes
Arabian mountains
Socotra
North-east Somalia
Central Ethiopian highlands
South Ethiopian highlands
Central Somalian coast
East Zairean lowlands
Albertine Rift Mountains
Kenyan mountains
Serengeti
Kenyan and Tanzanian coastal
forests
Eastern Arc Mountains
South Zambia
East Zimbabwean mountains
South-east African coast
South-east African grasslands
Cape region
Granite Seychelles
Aldabra
Comoro Islands
Mayotte
West Madagascan dry forest
East Madagascan humid forests
Central Madagascan lakes
West Madagascan coastal wetlands
South Madagascan Didiera scrub
Reunion
Mauritius
Rodrigues
Spain, Portugal 250-2,000
Cape Verde 0-160
Céte D'Ivoire, Ghana, 100-1,400
Guinea, Liberia, Sierra
Leone
Cameroon, Equatorial 700-2,900
Guinea, Nigeria
Cameroon, Gabon, 0-800
Equatorial Guinea, Nigeria
Sao Tome and Principe 0-1,000
Sao Tome and Principe 0-2,000
Angola 0-1,500
St Helena 0-300
USSR, Turkey 1,500-4,000
Cyprus 0-1,900
Iraq, Iran 0-100
Saudi Arabia, Yemen 1,800-3,200
Yemen 0-1,400
Somalia 300-2,100
Ethiopia 1,300-3,100
Ethiopia 1,275-2,300
Somalia 0-100
Uganda, Zaire 700-1,500
Burundi, Rwanda, Uganda, 1,000-3,200
Zaire
Kenya, Tanzania 1,100-3,700
Kenya, Tanzania 1,100-2,100
Kenya, Tanzania 0-500
Malawi, Mozambique, 750-3,000
Tanzania
Botswana, Zambia, 600-1,000
Zimbabwe
Mozambique, Zimbabwe 1,200-2,400
Mozambique, South Africa 0-100
Lesotho, South Africa 1,700-2,200
South Africa 0-1,000
Seychelles 0-900
Seychelles 0-8
Comoros 400-2,600
Comoros 0-1,700
Madagascar 0-800
Madagascar 0-2,300
Madagascar 750-1,500
Madagascar 0-100
Madagascar 0-200
Reunion 200-2,300
Mauritania 300-800
Reunion 0-390
187
Centres of Species Diversity
HABITAT(S)
forest, rocky
rocky, mixed
forest
forest
forest
forest,
mixed
forest
forest,
mixed
grassland,
mixed
rocky, mixed
forest, scrub
wetland,
mixed
scrub, mixed
scrub,
grassland
rocky, mixed
rocky, scrub
scrub, mixed
desert,
grassland
forest
forest
forest,
mixed
savanna
forest
forest
forest,
savanna
forest
forest, scrub
grassland
forest,
mixed
forest
forest
forest
forest
forest
forest
wetland
wetland,
forest
scrub, forest
forest
forest, scrub
forest, scrub
SIZE (km?)
8,100
4,000
113,000
7,300
40,000
140
860
14,000
200
64,000
9,300
40,000
59,000
3,500
41,000
37,000
15,000
1,200
49,000
44,000
46,000
47,000
8,800
39,000
47,000
4,900
43,000
60,000
24,000
240
160
1,900
360
30,000
112,000
2,000
5,000
30,000
2,500
1,900
100
SPP. R.
8.5
4.0
5.5
27.0
5:5
8.8
16.8
14.5
6.0
2.0
2.0
2.0
7.0
6.0
5.0
4.0
4.0
2.0
5.0
38.5
6.5
3.0
7.5
28.0
2.0
3.0
2.5
2.0
3.5
10.0
2.7
11.2
4.2
3.5
17.5
2.0
2.0
8.0
5.0
8.0
2.0
1. Biological Diversity
Table 15.7
West China
Western Himalayas
Indus valley
Western Ghats
Sri Lanka
Tibetan valleys
South Tibet
Eastern Himalayas
Assam plains
Tirap Frontier
Qinghai mountains
Central Sichuan mountains
West Sichuan mountains
South Chinese forests
Yunnan mountains
Burmese plains
Andaman Islands
Nicobar Islands
Annamese lowlands
Hainan
Da Lat Plateau
Cochinchina
Shanxi mountains
Fujian mountains
Taiwan
Nansei Shoto Islands
Ogasawara Islands
Endemic bird areas of the world
China
Afghanistan, India,
Nepal, Pakistan
India, Pakistan
India
Sri Lanka
China
China
Bhutan, China, India,
Myanmar, Nepal
Bangladesh, India
India, Myanmar
China
China
China
China
China, Myanmar
Myanmar
India
India
Laos, Viet Nam
China
Viet Nam
Viet Nam
China
China
Taiwan
Japan
Japan
SOUTH-EAST ASIAN ISLANDS AND AUSTRALIA
Luzon mountains
Luzon lowlands and foothills
Mindoro
Negros and Panay
Cebu
Palawan
Samar, Leyte, Bohol and Mindanao
lowlands
Mindanao mountains
Sulu Archipelago, excluding Basilan
Bornean mountains
Sumatra and Peninsular Malaysia
Enggano
Javan and Balinese mountains
Javan and Balinese lowlands
Flores and associated islands
Sumba
Timor and associated islands
Tanimbar and associated islands
Talaud and Sangir Islands
Sulawesi mountains
Sulawesi lowlands
Banggai and Sula Islands
Buru
Philippines
Philippines
Philippines
Philippines
Philippines
Philippines
Philippines
Philippines
Philippines
Indonesia, Malaysia
Indonesia
Indonesia
Indonesia
Indonesia
Indonesia
Indonesia
Indonesia
Indonesia
Indonesia
Indonesia
Indonesia
Indonesia
Indonesia
188
900-1,300
1,600-3,600
0-200
0-2,450
0-2,260
3,600-4,600
2,700-5,000
900-4,000
0-1,000
500-1,800
1,800-5,100
1,500-3,600
2,700-4,900
300-1,900
1,500-3,650
0-1,000
0-700
0-600
0-1,500
500-1,800
900-2,300
0-1,200
2,000-2,800
200-2,000
300-3,300
0-500
100-400
350-2,800
0-1,300
0-1,500
0-1,300
0-1,300
0-1,000
0-1,500
700-3,000
0-790
0-3,000
600-3,000
0-150
800-3,000
200-800
0-2,300
0-1,400
0-2,600
0-1,750
0-1,700
500-3,000
0-2,000
0-2,300
0-1,750
desert, scrub
forest
wetland,
scrub
forest
forest
scrub, rocky
scrub, forest
forest
wetland,
grassland
scrub, mixed
rocky, mixed
forest
forest, mixed
forest
forest
scrub,
agricultural
forest
forest
forest
forest
forest
forest
forest
forest
forest
forest, mixed
forest, mixed
forest
forest
forest
forest
forest
forest
forest
forest
forest
forest
forest
forest, mixed
forest
forest, scrub
forest
forest
forest
forest
forest
forest
forest
forest
forest
15,000
33,000
37,000
28,000
36,000
7,900
18,000
70,000
43,000
14,000
22,000
30,000
24,000
11,000
26,000
16,000
8,200
2,000
12,000
12,000
7,400
15,000
14,000
45,000
36,000
4,500
100
36,000
12,000
10,000
26,000
5,100
14,000
66,000
32,000
1,700
27,000
53,000
370
18,000
16,000
36,000
11,000
26,000
5,600
1,600
24,000
24,000
6,900
8,000
2.0
9.0
2.0
16.0
23.0
2.0
2.0
22.8
3.8
2.0
3.0
9.8
3.0
4.0
3.3
2.0
10.0
7.0
5.3
2.8
5.1
2.5
2.0
3.8
15.3
9.3
1.5
11.8
14.8
7.3
10.6
1.0
17.8
15.6
21.6
4.5
27.8
25.2
2.0
23.9
4.0
23.2
11.2
26.7
31.1
7.1
30.8
16.9
11.9
18.4
Table 15.7
NAME
Endemic bird areas of the world
POLITICAL UNIT(S)
SOUTH-EAST ASIAN ISLANDS AND AUSTRALIA (continued)
Seram
Halmahera
West Papuan Islands and Vogelkop
lowlands
Vogelkop mountains
Geelvink Bay Islands
North New Guinean mountains
North New Guinean lowlands
Adelbert and Huon mountains
Central New Guinean high mountains
Central New Guinean mid mountains
Trans-Fly and Upper Fly
Christmas Island
Kimberley and the Top End
Cape York
Atherton region
South-west Australia
Murray-Darling region and adjoining
coast
South-east Australia
Tasmania
NORTH AND CENTRAL AMERICA
California
Guadalupe Island
Baja California
Sierra Madre Occidental
North-west Mexican Pacific slope
Sierra Madre Oriental
North-east Mexican Gulf slope
Central Mexican marshes
Yucatan Peninsula
Revillagigedo Islands
Central Mexican highlands
Sierra Madre del Sur
Isthmus de Tehuantepec
North Mesoamerican highlands
North Mesoamerican Pacific slope
South Central American Caribbean
slope
South Central American Pacific slope
Costa Rican and Panamanian
highlands
North Choco and Darien lowlands
Darien highlands
Cocos Isles
Cuba and the Bahamas
Jamaica
Indonesia
Indonesia
Indonesia
Indonesia
Indonesia
Indonesia, Papua New
Guinea
Indonesia, Papua New
Guinea
Papua New Guinea
Indonesia, Papua New
Guinea
Indonesia, Papua New
Guinea
Indonesia, Papua New
Guinea
Christmas Island
Australia
Australia
Australia
Australia
Australia
Australia
Australia
USA
Mexico
Mexico
Mexico, USA
Mexico, USA
Mexico, USA
Mexico, USA
Mexico
Belize, Guatemala,
Honduras, Mexico
Mexico
Mexico
Mexico
Mexico
Belize,El Salvador,
Guatemala, Honduras,
Mexico, Nicaragua
Mexico, Guatemala, El
Salvador, Nicaragua,
Honduras
Costa Rica, Guatemala,
Nicaragua, Panama
Costa Rica, Panama
Costa Rica, Nicaragua,
Panama
Colombia, Costa Rica,
Panama
Colombia, Panama
Costa Rica
Bahamas, Cuba, Turks
and Caicos Is
Jamaica
189
ALTITUDE (m)
0-1,750
0-1,750
0-900
600-3,000
0-700
600-2,200
0-900
500-3,500
2,700-4,600
500-3,800
0-1,000
0-350
0-1,700
0-500
0-1,700
0-500
0-500
0-1,200
0-1,600
0-550
0-1,300
0-1,000
1,200-3,050
0-1,000
1,800-3,500
0-1,000
1,500-2,500
0-300
0-300
900-3,500
300-2,000
0-1,000
600-3,000
0-1,050
0-1,200
0-1,500
600-3,350
0-1,000
600-1,600
0-700
0-2,000
0-2,200
Centres of Species Diversity
HABITAT(S)
forest
forest
forest
forest
forest, mixed
forest
forest
forest
forest, mixed
forest
forest, wetland
forest
rocky, mixed
mixed
forest
mixed
scrub, mixed
forest
forest, mixed
forest, scrub
mixed
mixed
forest
forest, scrub
forest
mixed
wetland
forest, scrub
scrub, forest
scrub, forest
forest, scrub
scrub, forest
forest
forest, scrub
forest
forest
forest
forest
forest
forest, scrub
forest, scrub
forest, scrub
SIZE (km?)
14,000
29,000
14,000
26,000
3,200
11,000
32,000
19,000
6,800
98,000
64,000
140
105,000
43,000
28,000
115,000
98,000
85,000
68,000
30,000
280
17,000
36,000
14,000
16,000
77,000
10,000
138,000
280
41,000
18,000
7,700
68,000
15,000
25,000
24,000
27,000
14,000
4,200
47
93,000
11,000
SPP.R
19.3
32.5
11.6
13.3
9.6
4.5
7.1
8.3
13.5
32.8
6.6
2.0
13.0
4.3
14.8
13.5
5.5
9.5
15.0
3.0
2.0
3.0
3.5
9.0
2.0
4.0
2.0
15.1
5.0
15.8
6.8
2.3
21.0
3.0
8.0
13.0
52.5
10.0
13.5
3.0
21.3
30.3
1. Biological Diversity
Table 15.7
NAME
Endemic bird areas of the world
POLITICAL UNIT(S)
NORTH AND CENTRAL AMERICA (continued)
Hispaniola
Puerto Rico
East Caribbean
SOUTH AMERICA
North Choco and Darien lowlands
Darien highlands
Guianas
Tepuis
Cordillera de Caripe and Paria
Peninsula
North Venezuelan mountains
Venezuelan llanos
Merida mountains
Guajiran lowlands
Santa Marta Mountains
Nechi lowlands
Eastern Andes of Colombia
Upper Rio Negro white sand forests
Cauca valley
Magdalena valley
Choco
Western Andes of Colombia and
Ecuador
Galapagos Islands
Central Andes of Colombia and
Ecuador
Eastern Andes of Ecuador
Napo lowlands
Ecuadorian dry forests
North Peruvian cloudforests
Maranon valley
North-east Peruvian riverine forests
East cordilleran ridgetop forests
East Peruvian cordilleras
North Peruvian coast
Western Andes of Peru
Junin grasslands
Eastern Andes of Peru
South-east Peruvian lowlands
South-east Peruvian Andes
South Peruvian Pacific slope
Upper Bolivian yungas
Lower Bolivian yungas
Bolivian Andes
East Bolivian lowlands
North Argentinian Andes
Dominican Republic,
Haiti
Puerto Rico
Antigua and Barbuda,
Anguilla, Netherlands
Antilles, Barbados,
Dominica, Grenada,
Guadeloupe, St Kitts-
Nevis, St Lucia,
Martinique, Montserrat,
St Vincent and
Grenadines, British
Virgin Is, Virgin Is (US)
Colombia, Costa Rica,
Panama
Colombia, Panama
Brazil, French Guiana,
Guyana, Suriname
Brazil, Guyana,
Venezuela
Venezuela
Venezuela
Colombia, Venezuela
Venezuela
Colombia, Venezuela
Colombia
Colombia
Colombia, Venezuela
Colombia, Venezuela
Colombia
Colombia
Colombia, Ecuador
Colombia, Ecuador
Ecuador
Colombia, Ecuador
Colombia, Ecuador, Peru
Brazil, Ecuador, Peru
Ecuador, Peru
Ecuador, Peru
Peru
Peru, Ecuador
Ecuador, Peru
Peru
Ecuador, Peru
Peru
Peru
Peru
Brazil, Peru
Peru
Chile, Peru
Bolivia, Peru
Bolivia, Peru
Argentina, Bolivia, Peru
Bolivia, Brazil
Argentina
190
ALTITUDE (m)
0-3,000
0-1,200
0-1,500
0-1,000
600-1,600
0-1,100
500-2,800
700-2,500
750-2,400
0-1,100
750-4,000
0-600
750-4,600
0-1,500
900-5,200
100-500
600-2,700
200-2,700
0-1,200
500-3,300
0-1,300
2,100-5,200
400-2,000
100-600
0-2,000
1,500-3,200
200-2,400
100-450
1,000-2,400
1,900-3,700
0-500
1,800-4,300
3,700-5,000
700-1,600
100-400
2,500-4,300
0-3,000
1,800-3,700
700-2,400
1,400-4,600
200-750
2,000-4,000
HABITAT(S)
forest, scrub
forest, mixed
forest, scrub
forest
forest
forest
forest
forest
forest
savanna, mixed
forest
scrub, forest
forest
forest
forest, wetland
forest
forest
forest
forest
forest
scrub, forest
forest, mixed
forest
forest
forest, scrub
forest
forest, scrub
forest
forest
forest
scrub, mixed
scrub, forest
wetland,
grassland
forest
forest
forest, scrub
scrub, mixed
forest
forest
scrub, forest
forest, grassland
scrub, mixed
SIZE (km?)
76,000
9,000
6,600
14,000
4,200
174,000
35,000
4,000
7,100
57,000
18,000
36,000
5,400
28,000
67,000
10,000
19,000
29,000
59,000
27,000
8,000
37,000
24,000
129,000
57,000
9,200
11,000
11,000
8,900
44,000
31,000
59,000
17,000
11,000
155,000
13,000
76,000
19,000
38,000
32,000
93,000
17,000
SPP.R.
26.1
17.9
30.3
10.0
13.5
11.5
39.0
8.6
11.3
2.0
18.0
10.5
17.6
8.5
22.2
11.5
7.5
8.0
17.0
37.7
23.0
15.8
13.0
8.0
47.5
6.0
11.0
2.0
6.5
22.0
5.5
20.0
4.0
6.5
14.0
11.0
8.0
15.0
20.5
13.0
7.0
6.0
Table 15.7
NAME
SOUTH AMERICA (continued)
Argentinian grasslands
Argentinian cordilleras
Juan Fernandez Islands
Central Chile
Tierra del Fuego and the Falklands
Central Amazonian Brazil
West Amazonian Brazil
Fernando de Noronha
North-east Brazilian caatinga
Alagoan Atlantic slope
Bahian deciduous forests
Minas Gerais deciduous forests
Serra do Espinaco
Bahian and Espirito Santo Atlantic
slope
South-east Brazilian lowland to
foothills
South-east Brazilian mountains
South-east Brazilian Araucaria forest
Entre Rios wet grasslands
PACIFIC ISLANDS
Mariana Islands
Yap
Palau Islands
Micronesian Islands
Admiralty Islands
St Matthias Islands
New Britain and New Ireland
D’Entrecasteaux and Solomon Sea
Islands
Louisiade Archipelago
Solomon Islands
San Cristobal
Rennell Island
Vanuatu and the Santa Cruz Islands
New Caledonia and the Loyalty Islands
Samoan Islands
Fijian Islands
Norfolk Island
Lord Howe Island
New Caledonia North Island
South Island
Auckland Islands
New Caledonia Islands
Northwestern Hawaiian Islands
Hawaiian Islands
Hawaii
Marquesas Islands
Society Islands
Tuamotu Archipelago
Lower New Caledonia Islands
Pitcairn Islands
Endemic bird areas of the world
POLITICAL UNIT(S)
Argentina
Argentina
Chile
Argentina, Chile
Argentina, Chile,
Falklands
Brazil
Brazil
Brazil
Brazil
Brazil
Brazil
Brazil
Brazil
Brazil
Argentina, Brazil,
Paraguay
Brazil
Argentina, Brazil,
Paraguay
Argentina, Uruguay
Guam, N Marianas
Micronesia
Palau
Micronesia
Papua New Guinea
Papua New Guinea
Papua New Guinea
Papua New Guinea
Papua New Guinea
Papua New Guinea, Solomon
Is
Solomon Is
Solomon Is
Solomon Is, Vanuatu
New Caledonia
American Samoa, Samoa
Fiji
Australia
Australia
New Zealand
New Zealand
New Zealand
New Zealand
USA
USA
USA
French Polynesia
French Polynesia
French Polynesia
French Polynesia
Pitcairn
ALTITUDE (m)
100-500
1,600-2,900
0-1,300
0-1,600
0-1,200
0-300
0-400
te)
0-900
0-1,000
250-900
300-500
700-1,600
0-600
0-1,500
500-2,200
0-1,000
0-200
0-950
0-180
0-240
0-800
0-700
0-650
0-2,200
0-2,200
0-800
0-2,500
0-2,000
0-110
0-1,800
0-1,600
0-1,200
0-1,200
0-320
0-760 +
0-2,000
0-2,500
0-600 +
0-270+
0-300
0-3,100
0-3,100
0-1,200
0-1,700
0-110
0-700
0-33
Centres of Species Diversity
HABITAT(S)
scrub, wetland
grassland, mixed
forest, scrub
forest, scrub
grassland,
wetland
forest
forest
forest, scrub
forest, scrub
forest
forest
forest
grassland, scrub
forest
forest
forest
forest
wetland
forest, mixed
mixed, forest
forest, mixed
forest, mixed
forest, mixed
forest, mixed
forest, mixed
forest, mixed
forest
forest
forest
forest, mixed
forest, mixed
forest, mixed
forest, mixed
forest, mixed
forest, mixed
forest, scrub
forest
forest, mixed
grassland
mixed
scrub, mixed
forest
forest
forest
forest, mixed
forest, plantation
forest, mixed
forest
SIZE (km?)
34,000
10,000
180
137,000
119,000
35,000
30,000
26
100,000
30,000
8,000
10,000
30,000
40,000
50,000
65,000
30,000
25,000
1,000
56
500
580
1,900
300
46,000
3,400
1,300
32,000
3,300
840
16,000
19,000
3,000
18,000
35
6,200
10,000
1,000
400
690
190
36
SPP.R.
3.0
3.0
3.0
10.0
9.0
16.0
2.0
2.0
9.5
13.7
2.5
2.0
5.5
10.7
44.2
22.5
4.0
3.0
10.8
4.2
12.0
13.1
8.6
4.5
41.4
2.9
5.7
51.3
19.0
7.3
20.4
23.3
11.7
27.8
4.0
2.0
1.5
8.5
1.0
5.0
4.0
20.5
Wiles)
10.5
5.3
7.3
6.6
3.5
Note: Spp. R. is an index of the numbers of restricted-range species occurring in each EBA taking into account the sharing of species between
EBAs.
Table supplied by ICBP
191
1. Biological Diversity
16. SPECIES EXTINCTION
Species extinction is a natural process. The fossil record
suggests that all species have a finite lifespan and that the
vast majority of species that have ever existed are now
extinct, with extinct species outnumbering living species by
a factor of perhaps a thousand to one.
Species become extinct when all individuals die without
producing progeny. They disappear in a different sense
when a species lineage is transformed over evolutionary
time, or divides into two or more separate lineages (so-
called pseudo-extinction). The relative frequency of true
extinction and pseudo-extinction in evolutionary history is
unknown, although the former’s great importance is
demonstrated by the disappearance of entire, and once
highly diverse, lineages such as trilobites and ammonites.
HOW SPECIES BECOME VULNERABLE TO
EXTINCTION
Two broad categories of process are believed to affect the
dynamics of populations, and provide the fundamental
mechanisms of species extinction:
© deterministic processes (or cause and effect relationships)
e.g. glaciation or direct human interventions such as
deforestation
stochastic processes (chance or random events), which
may act independently or influence variation in
deterministic processes.
The magnitude of the effects of these processes depends on
the size and degree of genetic connectedness of populations.
Four types of stochastic processes can be distinguished
(Shaffer, 1987): demographic uncertainty (resulting from
random events in the survival and reproduction of
individuals); environmental uncertainty (due to
unpredictable changes in weather, food supply, disease, and
the populations of competitors, predators, or parasites);
natural catastrophes (floods, fires or droughts); and genetic
uncertainty (random changes in genetic make-up, to which
several factors contribute).
Models of the effects of stochastic processes suggest that:
e demographic uncertainty is only a hazard for relatively
small populations (numbering tens or hundreds of
individuals)
there is no critical population size that once reached
guarantees a high level of long-term security from
environmental uncertainty
progressively larger increases in population size yield
diminishing returns in persistence times for a given
catastrophic event.
When demographic and environmental uncertainty interact,
their effects compound each other, so that in a variable
environment any loss in population size proportionally
increases the chance of population extinction. Thus, to be
reasonably certain of conserving a species for a significant
length of time, one must preserve either very large
population sizes (hundreds to millions of individuals or
more, depending on the biology of the species) or numerous
populations (Schaffer, 1987).
192
The isolation of populations
The ‘equilibrium theory’ of island biogeography developed
by MacArthur and Wilson (1963 and 1967) is an extension
of the species-area relationship (see Chapter 5). Whilst
originally used to model species richness and turnover on
real islands, it has subsequently been used to predict
changes in species number in isolated habitat islands.
The area of an island sets an upper limit to the maximum
population size of each species. Since small populations are
inherently more prone to extinction than large (for reasons
discussed above), extinction rates tend to be inversely
proportional to island area. Successful colonisation by new
species is not affected so much by area as by the degree of
isolation of the island: islands near to the mainland or to
other islands are colonised at higher rates than those farther
away. Increased isolation of populations not only reduces
the incidence of colonisation by new species, but decreases
the probability that immigrants of an existing species will
arrive. Over time, an equilibrium is eventually reached on
any island at which the loss of species through extinction is
balanced by the arrival and colonisation of new species.
A later modification of the theory incorporates the ‘rescue
effect’ (Brown and Kodric-Brown, 1977). The immigration
of new, unrelated individuals can play an important role in
maintaining an isolated population, because their
demographic and genetic contributions tend to increase its
size and genetic fitness, thereby reducing the possibility that
it will become extinct. The significance of the rescue effect
is that fewer immigrants are needed to rescue an existing
population than to successfully found a new one.
Island biogeographic theory has far-reaching implications
for conservation biology. Rates of habitat modification are
currently so high that virtually all natural terrestrial habitats
and protected areas are destined to become ecological
‘islands’ in surrounding ‘oceans’ of habitat much altered by
human activity. Not only is the total area of many natural
habitats rapidly decreasing, but those large natural habitat
islands that now exist are being fragmented into
archipelagos of habitat islands. This process of
fragmentation and isolation is predicted to lead directly and
indirectly to accelerated species extinctions at both the local
and global scales.
Consequences of insularisation
The combination of short- and long-term insularisation
effects is predicted to reduce the number of species to a
lower equilibrium. A study of understorey birds in
fragments of tropical forest ranging from 0.1 to 571ha in
the Usambara Mountains, Tanzania, found just this result
(Newmark, 1991). Since separation, smaller forest
fragments have lost more bird species than larger areas, and
more isolated fragments have lost more species than those
close to a source of potential colonists. Similarly, Klein
(1989) observed communities of dung and carrion beetles
(subfamily Scarabaeinae) in fragmented habitat patches of
different sizes in the Amazon rain forest of Brazil. He
found that forest fragments had lower species richness, an
(1989) observed communities of dung and carrion beetles
(subfamily Scarabaeinae) in fragmented habitat patches of
different sizes in the Amazon rain forest of Brazil. He
found that forest fragments had lower species richness, an
increased proportion of rare species, and _ sparser
populations in comparison with continuous undisturbed
forest. These differences were more pronounced in small
fragments (< lha) than large.
Many researchers, however, are now convinced that
calculation of rates of species loss in habitat islands or
reserves using the species-area relationship is unjustified as
a basis for detailed conservation recommendations.
Boeckeln and Gotelli (1984) argue that the models
developed ignore species identity, habitat heterogeneity and
population sizes, and have such wide margins of error that
they have low explanatory power and give unreliable
estimates. For example, Soulé et al. (1979) predicted on the
basis of a simple species-area model that the Serengeti
National Park will lose 50% of its large mammals (some 15
ungulate species) in the first 250 years of isolation, while
Western and Ssemakula (1981) attempted to incorporate
habitat diversity data and predicted that only one species
will be lost. Zimmerman and Bierregard (1986) argue that
beyond the ecological truism that species richness increases
with area, the equilibrium theory of biogeography has
revealed little that is of "real value for planning real
reserves in real places". In designing reserves to protect
Central Amazonian forest frogs, Zimmerman and
Bierregard consider that critical breeding habitat and places
that contain quality habitat at high density must be found
before the reserve size question is addressed. In general,
biologists need empirical studies that directly measure the
effects of habitat fragmentation on specific groups (Klein,
1989).
Ecological correlates of vulnerability to extinction
There is considerable evidence that the number of species
in an isolated habitat will decrease over time, although the
probable rates of such extinctions (and whether the
equilibrium theory of island biogeography can be used to
predict these) are in dispute. The crucial issue for
conservationists now is whether those species which are
most at risk from extinction following habitat fragmentation
can be predicted from a knowledge of their biology and
ecology. At least nine ecological or life history traits (some
of which may actually be highly correlated with each other)
have been proposed as factors determining an animal
species sensitivity to fragmentation (Karr, 1991; Laurance,
1991):
Rarity
Several studies have found that the abundance of a species
prior to habitat fragmentation is a significant predictor of
extinction. For example, Newmark (1991) found that after
fragmentation, rare understorey bird species occupied fewer
forest fragments per species than common ones. This is
only to be expected, since fewer individuals of a rare
species than a common species are likely to occur in habitat
fragments, and the mechanisms of extinction mean that
small populations are inherently more likely to become
extinct than large.
193
Species Extinction
Dispersal ability
If animals are capable of migrating between fragments or
between ‘mainland’ areas and fragments, the effects of
small population size may be partly or even greatly
mitigated by the arrival of ‘rescuers’. Species that are good
dispersers may therefore be less prone to extinction in
fragmented habitats than poor dispersers.
Degree of specialisation
Ecological specialists often exploit resources which are
patchily distributed in space and time, and therefore tend to
be rare. Specialists may also be vulnerable to successional
changes in fragments and to the collapse of coevolved
mutualisms or food webs.
Niche location
Species adapted to, or able to tolerate, conditions at the
interface between different types of habitats may be less
affected by fragmentation than others. For example, forest
edge species may actually benefit from habitat
fragmentation.
Population variability
Species with relatively stable populations are less vulnerable
than species with pronounced population fluctuations, since
they are less likely to decline below some critical threshold
from which recovery becomes unlikely.
Trophic status
Animals occupying high trophic levels usually have small
populations: e.g. insectivores are far fewer in number than
their insect prey and, as noted above, rarer species are
more vulnerable to extinction.
Adult survival rate
Species with naturally low adult survival rates may be more
likely to become extinct, as Karr (1991) has proposed for
island birds on Barro Colarado Island, Panama.
Longevity
Long-lived animals are less vulnerable to extinction than
short-lived.
Intrinsic rate of population increase
Populations which can expand rapidly are more likely to
recover after population declines than those which cannot.
Laurance (1991) has, however, studied extinction proneness
among 16 species of non-flying land mammals in
fragmented rain forest in Queensland, Australia. Seven
traits were examined: body size, longevity, fecundity,
trophic level, dietary specialisation, natural abundance in
continuous rain forest, and ‘matrix’ abundance (abundance
of the species in modified habitats surrounding original
fragments). Of these, matrix abundance was the best
predictor of vulnerability. Once its effects were removed,
partial correlations showed no other significant predictors
of extinction proneness.
Laurance therefore suggests that tolerance of modified
habitats is important in determining survival in fragmented
habitats. Species that were able to exploit modified habitats
tended to remain stable or even to increase in number in
1. Biological Diversity
fragments, whereas those that avoided these habitats tended
to disappear. The most vulnerable species even avoided
using the corridors of secondary growth forest that existed
along streams, a finding which highlights the importance of
maintaining corridors of primary vegetation to act as
pathways for dispersing individuals between patches of
habitat.
Viable populations, genetic variation and extinction
Increasingly the attention of conservationists has become
focused on the management and preservation of isolated
small populations confined to habitat islands, usually in
protected areas. An essential requirement is to ascertain
how many individuals of a species should be conserved in
order to ensure its survival in a particular area. There are
two approaches to estimating such a Minimum Viable
Population (MVP) size, the demographic and the genetic.
The process of applying a demographic or genetic MVP
model to a particular species or population, and proposing
the management interventions that should be undertaken to
increase its chances of survival, is known as Population
Viability Analysis (PVA).
In the demographic approach, estimates of a population’s
average growth rate (which is in part determined by the
species’s body size), the variance in this growth rate
attributable to environmental fluctuations, and _ the
population’s maximum size, are used in mathematical
models to calculate its expected persistence time to
extinction. There are two main factors that need to be
considered: the population size, and the length of time it
requires to be preserved. Normally, an MVP is taken to be
that size of population that has a 95% probability of
persistence for x number of years, where for consistency x
is usually taken as either 100 or 1,000.
Clearly, there is no such thing as a standard MVP that can
be applied to all species. Belovsky (1987) has calculated
that over a range of body masses from 10g (the size of a
European Common Shrew Sorex araneus) to 10°g (the size
of a Black Rhinoceros Diceros bicornis), MVP sizes for
mammals range from hundreds to millions. The Minimum
Area Required (MAR) to support these populations ranges
from tens to millions of square kilometres. As body mass
increases, MVP size decreases, but larger mammals require
proportionately larger ranges. MARs are larger for
carnivores than for herbivores and larger for tropical than
for temperate species.
MVPs can also be examined from a genetic perspective, in
which not only the number of individuals surviving but their
genetic variation or heterozygosity are considered
important. In the long term, this genetic variation is
necessary for evolution by natural selection to occur, and is
required for adaptation to potential future changes in the
environment. In the short term, heterozygosity is positively
correlated with fitness, including survival, disease
resistance, growth and developmental rate and stability
(Allendorf and Leary, 1986).
Genetic variation can quickly be lost through breeding with
closely-related individuals (inbreeding) which leads to low
levels of heterozygosity and lowered offspring fitness, a
194
phenomenon known as inbreeding depression (Falconer,
1981). The most likely explanation is that new mutations,
which are almost always harmful, can accumulate in a
species genome providing they are fully or partially
recessive and are not therefore expressed. Inbreeding
increases the probability that the effects of these harmful
genes will be expressed.
Franklin (1980) has proposed that in the short term an
effective population size of 50 is the MVP required to
guard against the negative effects of inbreeding for a
population of large mammals with no immigration or
introduction of unrelated stock. Populations of this size will
nevertheless eventually become inbred over time, to a
degree directly related to the generation interval (randomly-
breeding populations of 50 mice will become more inbred
in a decade than 50 elephants will in a century). In the long
term an effective population size of 500, corresponding to
a real population size of several times this number, has
been suggested as a suitable genetic MVP for large
mammals, since in a population of this size rates of
mutation will renew genetic variation as quickly as it is lost
by inbreeding and genetic drift (Franklin, 1980; Lande and
Barrowclough, 1987).
Although biologists have suggested the figures quoted above
as useful first estimates of MVP sizes, in both the
demographic and the genetic approach the actual numerical
value arrived at depends not only on the criteria chosen to
define the MVP (e.g. the number of years the population is
required to persist) but also on the values of the parameters
used in the model. These values cannot always be assigned
in an objective manner. Thus even for one particular
species there is no single number that is universally valid,
and this reservation is doubly true when different species
are compared. Each situation is unique and should be
considered separately. For example, a species that exhibits
a boom and crash population cycle will require a larger
MVP than one which inhabits a stable environment and
whose population is relatively stable.
Both MVPs and PVAs have now been applied to a variety
of species. Examples include: large mammals such as the
Sumatran Rhinoceros Didermocerus sumatrensis and the
Florida Panther Felis concolor coryi; and birds such as the
Bali Starling Leucopsar rothschildi, Caribbean parrot
species, and Asian Hornbills.
Analysis of the existing worldwide protected areas system
indicates that few if any large mammal species will be
adequately conserved with the current scale of ecosystem
coverage, as most protected populations are too small to
constitute MVPs (Grumbine, 1990).
The fact that a population has declined in number to below
the theoretically determined MVP does not automatically
mean that its situation should be considered hopeless. Some
species, such as the Northern Elephant Seal Mirounga
angustirostris (Bonner and Selander, 1974) and captive
populations of Golden Hamster Mesocricetus auratus have
survived through population bottlenecks of just a few
individuals, following which numbers have increased to
substantial levels. Eventually, if a large population is re-
established, genetic variation may be regenerated by
mutation, thus restoring the potential for adaptive evolution.
These examples, however, may be the exceptions rather
than the rule. Other species that have declined to such low
levels may have vanished altogether. Even if populations do
recover numerically from a bottleneck, inbreeding and
consequent loss of heterozygosity may cause noticeable
declines in fitness effectively prejudicing the species long-
term chances of survival. For example, O’Brien ef al.
(1985) found high rates of juvenile mortality, incidence of
sperm abnormalities, and susceptibility to disease in several
populations of Cheetah Acinonyx jubatus, and attributed this
to the low level of genetic variation found in all Cheetah
populations examined.
Perhaps the most compelling evidence to date of the
negative consequences of population bottlenecks comes from
a study of Lion Panthera leo in Ngorongoro Crater and the
neighbouring Serengeti Plains in Tanzania. In 1962 the
relatively isolated Lion population in the Crater dropped
from around 70 individuals to 10 as a result of an outbreak
of biting flies Stomoxys calcitrans. The population has since
recovered to its pre-plague levels. Packer et al. (1991) have
found that compared to the larger outbred population of
Serengeti Lions, those in Ngorongoro suffer high levels of
sperm abnormality. Their reproductive performance has
also diminished over the years since the bottleneck, and
both effects are apparently correlated with the lower levels
of heterozygosity in the Ngorongoro population.
Metapopulation theory
The MVP models discussed so far have considered all
individuals as belonging to a single isolated population,
which is rarely the case in the real world. In practice most
species are patchily distributed, and are best regarded as a
population of subpopulations, or a metapopulation, in which
subpopulations are geographically isolated but
interconnected by patterns of gene flow, extinction and
recolonisation. Thus, studies over a 25-year period by
Erhlich and colleagues of a purported single population of
Checkerspot Butterfly Euphydryas editha bayensis in the
Jasper Ridge Preserve (USA) demonstrated that although
the population occupied three nearly contiguous habitat
patches, it actually consisted of three demographic units
whose sizes fluctuated independently in response to annual
changes in rainfall. One of these units became extinct, was
re-established by immigration, and became extinct again
several years later (Wilcox and Murphy, 1985). Relaxing
the single population assumption of the MVP model so that
immigrants can be received from neighbouring populations
will lengthen the projected persistence times.
Habitat heterogeneity and the existence of many
subpopulations are an important element of population
dynamics, and have profound implications for conservation
biology. Pulliam (1988) introduced a simple model of
metapopulation dynamics incorporating density-dependent
immigration as the linking factor between source and sink
populations in severely fragmented habitats. In his model,
a limited number of reproductively successful ‘source’
subpopulations produce an excess of offspring over and
above the number that the habitat can absorb. The surplus
individuals migrate to other less favourable areas, occupied
195
Species Extinction
by ‘sink’ subpopulations which would be doomed to
extinction without persistent immigration.
Supporting evidence for the source-sink metapopulation
theory is available from a number of field studies; for
example, King and Mewaldt (1987) found that isolated
montane populations of White-crowned Sparrows
Zonotrichia albicollis were unable to persist without
periodic immigration.
Metapopulation theory should help biologists determine
which populations are priorities for conservation. The
importance of identifying and preserving source populations
and habitats is obvious: without them the metapopulation
cannot persist. However, the presence of breeding
individuals at a particular site does not necessarily indicate
that it is suitable for the species in the long term, since it
could still be a sink habitat. In general, source populations
will not only have higher annual reproduction rates than
annual mortality rates but will also have more stable
populations than sink populations. In the case of long-lived
species the identification of source populations will
therefore necessitate continuous, long-term monitoring. In
addition to the identification and protection of demographic
source populations, the conservation of buffer habitats and
marginal subpopulations should also be a part of
comprehensive conservation plans, and the long-term status
of even apparently secure metapopulations should be
carefully monitored.
Conclusion
Current models of the extinction process and estimates of
habitat loss, principally tropical forest, predict that species
extinctions are occurring at very high rates on both a local
and global scale. The primary cause is habitat modification
and fragmentation by human activities. This process not
only decreases overall population sizes of many species but
splits previously continuous populations into smaller isolated
sub-populations. Deterministic and stochastic effects mean
that small populations are more susceptible to extinction
than large. Conservation biologists have enlisted the help of
various theories and models to try and predict how many
species, and which ones, will be lost. It is possible to make
reasonable predictions of which species will be most
adversely affected by habitat fragmentation.
The species-area relationship is not now thought to be a
good predictor of species loss in habitat fragments, but has
implications for the design and positioning of reserves.
With a realisation that ecosystems are often best preserved
by concentrating on keystone species, efforts have switched
to conducting population viability analyses for selected
species in an attempt to estimate the minimum viable
population sizes that must be conserved to ensure their
long-term survival. MVPs can be examined from either the
demographic or genetic perspective - both approaches give
estimates of a similar order of magnitude. A shortcoming
of MVP estimates is that they consider only a single
population. The incorporation of metapopulation theory
should improve the accuracy and utility of these models,
and allow the identification of the most important
1. Biological Diversity
subpopulations, facilitating the determination of
conservation priorities.
A BRIEF HISTORY OF EXTINCTIONS
Knowledge of extinction patterns through geological time is
based on analysis of the fossil record, which represents a
small and highly biased sample of the taxa that have existed
- it may represent only one in every 20,000 species that has
existed. The best preserved group consists of marine
animals, chiefly invertebrates, with durable, highly
mineralised exoskeletons. Caution has to be exercised in
extrapolating from this group to others, particularly plants,
as they may show different patterns of extinction.
Mass extinction events in marine organisms
The fossil record indicates that overall extinction rates have
not been constant over time (Fig. 16.1). Around 60% of
extinctions have occurred in a number of relatively short
episodes. The earliest period for which there is evidence of
a major loss of diversity is during the late Precambrian,
around 700 Mya (million years ago) although the
Precambrian fossil record is too incomplete to allow
detailed analysis.
The fossil record for the Phanerozoic (i.e. from the
Cambrian to the present, see Fig. 16.2) is much more
detailed. During this time there have been five major ‘mass
extinction events’. These events took place late in each of
the Ordovician, Devonian, Permian, Triassic and
Cretaceous periods. By far the most severe was in the late
Permian (245 Mya). At that time, the number of families of
marine animals recorded in the fossil record declined by
54% and the number of genera by 78-84%. Extrapolation
from these figures indicates that species diversity may have
dropped by as much as 96%. The second most severe mass
extinction, at the end of the Ordovician (440 Mya), resulted
in the loss of 22% of families of marine taxa, a slightly
Extinction events in marine
organisms
Figure 16.1
Diversity 00
Bs 8 6 8 8 8
Source: Modified from Erwin, D.H., Valentine, J.W. and Sepkoski,
J.J. 1987. A comparative study of diversification events: the early
Palaeozoic versus the Mesozoic. Evolution 41(6).
Note: The curve plots diversity of marine animal families
and indicates five major extinction phases.
196
Figure 16.2 The geological time scale
Era Per iod Millions of
years ago
Quaternary 2
=
0)
3
N Tertiary
fe}
£ 66
Cretaceous
5
i) 138
(e)
N
° Jurassic
fa)
195
Triassic
245
Permian
290
Carboniferous
345
Devonian
UD
Zl 400
o
a Silurian
5 440
0 os
Ordovician
500
Cambr ian
580
Note: Dates are approximate; scale covers the Phanerozoic only.
greater figure than the late Devonian and late Triassic
events (21% and 20% respectively). The late Cretaceous
event was the least important, resulting in the loss of
around 15% of marine families.
The causes and timespans of these events have been the
subject of much debate and study. It is now widely accepted
that the late Permian mass extinction was a long-term event,
lasting for 5-8 million years. It appears to have been
associated with geologically-rapid global physical changes
(including the formation of the supercontinent Pangea),
climate change, and extensive, tectonically-induced marine
transgression and increased volcanic activity. There is no
direct evidence of a single, catastrophic event such as
impact by an extra-terrestrial body, although this cannot be
ruled out as a contributory factor in the event. Interpretation
of the late Triassic event is hampered by the absence of a
good stratigraphic record; some indications suggest this was
also a protracted period of extinction, although this is
uncertain. The late Devonian extinction also appears to have
spanned a considerable length of time, with elevated
extinction rates throughout much of the middle and late
Devonian. However, this extinction phase probably
consisted of a series of discrete shorter extinction events
rather than one protracted episode.
In contrast to these, the late Ordovician and late Cretaceous
extinctions are thought to have taken place over a much
shorter period. The late Ordovician event appears to be
correlated with global glaciation 439 Mya (the Hirnantian
glaciation) with three separate episodes of extinction spread
over only 500,000 years.
The late Cretaceous extinction is probably the best known,
but in terms of overall loss of diversity is also the least
important. There is some evidence that this extinction event
was associated with an extra-terrestrial impact, although this
remains controversial.
As well as these major mass extinction events, a large
number of less dramatic, but still significant, episodes can
be identified from the marine fossil record. It has been
argued that those following the late Permian extinction
event have a periodicity of 26-28 million years, indicating
some underlying unifying cause, although this remains
unproven. It is notable that these more minor events
account in total for more extinctions than the five major
events outlined above.
Mass extinctions in vertebrates
The vertebrate fossil record, especially for terrestrial
tetrapods, is much less amenable to analysis of extinction
rates than the invertebrate record chiefly because it is less
complete and less diverse. However, studies indicate that
tetrapods have been subject to at least six mass extinction
events since their appearance in the late Devonian, while
fishes have experienced eight such events since their
recorded origin in the Silurian. Some of these events
coincide with each other and with those recorded for marine
invertebrates; in particular, the five major mass extinction
events outlined above are paralleled by losses in vertebrate
diversity. The most significant is the late Permian event,
which is the largest recorded extinction both for fishes
(44% of families disappearing from the fossil record) and
tetrapods (58% of families disappearing). The late
Cretaceous event was more significant for tetrapods than for
other groups, with 36 of the 89 families in the fossil record
disappearing at this time. These families were, however,
Virtually confined to three major groups which suffered
complete extirpation - the dinosaurs, plesiosaurs and
pterosaurs. Most other major vertebrate taxa were almost
completely unaffected.
Evidence for correlation between the more minor extinction
events in vertebrates and the postulated periodic extinctions
in marine invertebrates is currently poor.
197
Species Extinction
Extinctions in vascular plants
In general, the plant fossil record does not clearly show the
same sudden mass-extinction events seen in the animal
record. Part of the explanation for this may lie in the nature
of the plant fossil record itself and in the difficulties in
interpreting it, but there also seem likely to be genuine
differences between plants and animals in patterns of
species origination and extinction. Plant extinction rates
(based on analysis of families and genera) do vary with
time, but in general, periods of elevated plant extinction
appear to be more protracted than animal extinction events
and do not usually coincide with them. It is argued that
these periods may be more to do with competitive
displacement by more developed plant forms, or with
gradual climatic change, than with any sudden catastrophic
events (Knoll, 1984).
The major exception to this is the end-Cretaceous
catastrophe, which appears to have had a major influence
on the structure and composition of terrestrial vegetation
and on the survival of species. Data from fossil leaves
suggest that perhaps 75% of late Cretaceous species became
extinct, although data from fossil pollens indicate a lower
though still significant level of extinction. During the
Tertiary there are two other periods of widespread enhanced
extinction rates, during the late Eocene and from the late
Miocene to the Quaternary, although in the latter, extinction
of taxa at generic level and above appears to have been
mainly regional rather than global.
Background extinction rates
A corollary of the finding that the majority of extinctions
recorded in the fossil record have taken place over
relatively short time periods (geologically speaking) is that
extinction rates for the remainder of the Phanerozoic have
been low.
The average lifespan of species in the fossil record is
around four million years which would give, at a very gross
estimate, a background extinction rate of four species each
year out of a total number of species of around 10 million.
However, it can be argued that the fossil record is heavily
biased towards successful, often geographically wide-
Tanging, species which undoubtedly have a far longer than
average persistence time. Most species will therefore
survive for less than four million years, and real extinction
rates at any given time will be correspondingly higher.
Nevertheless, even if background extinction rates were ten
times higher than this, extinctions amongst the 4,000 or so
living mammals would be expected to occur at a rate of
around one every 400 years, and amongst birds at one
every 200 years.
It is indisputable that the extinction rate in recent times has
been far higher than this and that man has been the
overwhelming cause. It is also widely accepted that
mankind is in danger of precipitating further extinctions on
a scale and at a rate at least comparable with those of the
major extinction events in the distant past.
1. Biological Diversity
Extinctions and the spread of mankind
Documenting man’s impact on the world’s biota, and in
particular quantifying species extinctions induced by man,
is difficult for a variety of reasons, associated with:
identifying species, especially those known only from sub-
fossil or fossil remains; unequivocally demonstrating that
extinction has occurred; and establishing a causal link
between man’s activities and extinction of the species in
question.
Man may have first had a significant impact on the survival
of other species during the late Pleistocene. Humans spread
into Europe and Asia about one million years ago but slow
advances in culture and technology seem to have restricted
the impact on the fauna of these regions. However, man’s
arrival on previously isolated continents, around 50,000
years ago in the case of Australia and 11,000 years ago for
North and South America, seems to coincide with large-
scale extinctions in certain taxa. The exact timings are
unclear and hence the cause and effect in each case are
open to debate. However, Australia lost nearly all its
species of very large mammals, giant snakes and reptiles,
and nearly half its large flightless birds around this time.
Similarly, North America lost 73 % and South America 80%
of their genera of large mammals at around the time of the
arrival of the first humans. In these cases there is more
direct evidence to link the events, although climatic
upheavals at around the same time could also be implicated.
EXTINCTIONS IN RECENT HISTORY
The European Age of Expansion in the 15th and 16th
centuries initiated another wave of extinctions. Indeed it has
often been assumed that all, or at least the great majority,
of modern man-induced extinctions date from this period.
However, this may well be based more on the fact that a
dramatic increase in documentation of natural phenomena,
in large measure induced by the great voyages of discovery,
also dates from this time.
It is now known that in some parts of the world a
significant number of extinctions occurred before the arrival
of Europeans. The Polynesians, who colonised the
Hawaiian Islands in the 4th and Sth centuries AD, appear
to have been responsible for exterminating around 50 of the
100 or so species of endemic land birds in the period
between their arrival and that of the Europeans in the late
18th century. A similar impact seems to have been felt in
New Zealand, which was colonised some 500 years later
than Hawaii. Here an entire avian megafauna, consisting of
members of the family Anomalopterygidae (the Moas) was
apparently exterminated, also by the end of the 18th
century. As with the late Pleistocene extinctions, there has
been some controversy over the extent to which humans
were responsible; however there is now a broad consensus
that man was indeed responsible, probably through a
combination of direct hunting and large-scale habitat
destruction through burning.
Although most information from this period relates to avian
extinctions, there is evidence that other groups, particularly
mammals, had been similarly affected. On Madagascar, in
addition to 6-12 ratites, including the Giant Elephantbird
198
Aepyornis maximus (the largest bird ever recorded), at least
14 lemur species, most of them larger than any surviving
species, have become extinct within the last 1,500 years, as
have two giant tortoises. In the Caribbean, at least two
ground sloths in the family Megalonychidae, several large
rodents and three insectivores in the family Nesophontidae
survived into the period of Amerindian settlement, but had
become extinct before Europeans arrived at the end of the
15th century. The case for man being solely responsible for
these extinctions is more equivocal than it is for New
Zealand. However, on balance this appears to remain the
most likely explanation, although it is possible that, on
Madagascar at least, climate change leading to progressive
desiccation of the environment also played a part.
While documentation has improved considerably since the
15th and 16th centuries, it still remains far from complete.
This applies even to the best known groups, namely birds
and mammals; for most lower vertebrates and virtually all
invertebrates knowledge of extinction rates remains
extremely scanty.
The main problem for documentation is that the majority of
the world’s species, especially tropical invertebrates, have
not been scientifically named. A significant percentage of
these may well become extinct before they have ever been
collected and described. Of described taxa, numbering
around 1.1 million animal species and around 270,000
vascular plants, accurate information on status and
abundance is available for only a tiny proportion. The vast
majority of the world’s species, even in the best-known
groups such as mammals and birds, are not subject to
systematic monitoring and species may be locally or
completely extirpated before their plight becomes known.
In general, it can only be stated with any confidence that a
taxon is extinct when unsuccessful attempts have been made
to locate it, or when it has not been sighted for several
decades. Animal species thought to have become extinct,
using this criterion and expert opinion, are listed in Table
16.1. Even here it is often difficult to demonstrate
unequivocally that a species has become extinct and
consequently several species are marked as possibly still
being extant. Many species may persist unrecorded (albeit
often in very low numbers) despite intensive efforts to
locate them. This is borne out by the periodic reappearance
of ‘Lazarus taxa’, after many years or decades of presumed
extinction. Plants (Table 16.2), some of which produce
seeds that can lie dormant and undetected for many years
before germination, present particular monitoring problems.
Historical records of extinctions may thus be expected to be
heavily biased, both taxonomically and geographically.
Taxonomically, informationon snails, particularly terrestrial
species, birds and mammals is good, while that for most
other groups is poor. Geographically, information on
Europe and North America (including Hawaii) is much
better than that for the rest of the world, although relatively
few species extinctions have been recorded in Europe in
recent times. Figures 16.6-16.10, taken from Table 16.6,
illustrate these biases.
These biases make analysis of extinction patterns
problematic. However, certain generalised patterns do
Table 16.1
MOLLUSCS BIRDS
ISLANDS 151 104
% of islands total 41.6 28.7
% of grand total 31.2 21.5
CONTINENTS 40 11
% of continents total 33.1 94
% of grand total 8.3 2.3
TOTALS 191 115
% of total on islands 79 90.4
% of TOTALS 39.5 23.8
Species Extinction
Summaries of animal extinctions on islands and continents
MAMMALS OTHER TOTAL
34 74 363
9.4 20.4 100
7 15.3 75
24 46 121
20 38 100
5 9.5 25
58 120 484
59 61.7 75
12 24.8 100
Note: these summaries do not take into account 4 species (2 birds, 1 mammal and one ‘other’) which are not assignable to either island or continent.
emerge. The most important of these is the preponderance
of extinctions on islands over those in continental areas
(Table 16.1). Exactly 75% of recorded animal extinctions
since 1600 have been on islands. For the three groups with
best information, the proportion of island extinctions varies
from 90% for birds to 58% for mammals, with molluscs
intermediate at 80%. Of the continental extinctions, at least
66% can be classified as aquatic species (this includes
amphibians and insects with aquatic larval stages but
excludes birds such as ducks and grebes). Most striking,
perhaps, is the very small number of extinctions recorded
to date in continental tropical forest ecosystems, which are
precisely the areas where mass extinction phenomena are
predicted to be taking place at present (see below).
There appear to be several reasons for the elevated
extinction level amongst island species. Most
straightforwardly, island species, especially those confined
to single islands, tend to have very restricted and
completely circumscribed ranges: they consist effectively of
single populations. Adverse factors are thus likely to affect
the entire species and bring about its extinction. In contrast,
continental species tend to occupy larger ranges existing as
meta-populations, with a number of more-or-less isolated
subpopulations. It is likely that some of these
subpopulations will not be affected by a given adverse
factor. Thus, the species itself will survive even if a number
of subpopulations are extirpated. These concepts apply both
to real islands and ecological islands, that is, areas of
habitat separated from other such areas by inhospitable
environments which act as an effective barrier to dispersal.
In this context, aquatic species in isolated inland waters
behave similarly to terrestrial species on isolated oceanic
islands, which helps to explain the significantly elevated
number of extinctions amongst continental freshwater
species.
Many island species are innately vulnerable to extinction
because of their biology. Species on islands have often
evolved in the absence of terrestrial predators and may
therefore be highly vulnerable to introduced predators.
Tameness, flightlessness and reduced reproductive rates
characterise many island birds and appear to have been
major contributory factors in their extinction, through
predation by humans or introduced species. Similarly, many
island land snails, such as the Hawaiian Achatinella and
French Polynesian Partula species, have low reproductive
rates and, apparently, no defences against introduced
199
predators, most notably the snail Euglandina. The elevated
species extinction rates on islands can also be ascribed to
taxonomic practices, as there has been a tendency for island
populations to be designated as full species when they may
more reasonably be regarded as subspecies of species on
adjacent islands or on the mainland.
Causes of Extinction
A brief analysis of the ‘Possible causes’ column of Table
16.4 shows that introduced animals and direct habitat
destruction by man have been major factors involved in
these extinctions, being implicated in 17% and 16%
respectively (see Fig. 16.3). These are equivalent to 39%
and 36% if only those extinctions for which causes are
assigned are counted. Hunting and deliberate extermination
also contribute significantly (23% of extinctions with known
cause). For a large number of animais, no information on
cause of extinction was known.
Figure 16.3 Causes of animal extinction
No cause assigned
Introduced
animals
Hunting
“Habitat destruction
id -)
Notes: These figures were compiled by giving each species a score of
1 in the appropriate category if there was only one cause, 0.5 in each
for two, etc. Where there were multiple causes C/D was counted as
one part, C and D as two parts.
Time Series
Figures 16.4 and 16.5 and Table 16.2 present a breakdown
of recorded extinctions in 30-year intervals from the year
1. Biological Diversity
Table 16.2 Time series of animal extinctions on islands and continents
ISLANDS
MOLLUSCS BIRDS MAMMALS OTHER TOTAL
TOTALS 151 104 Ka 74 303
1600-1629 i?) 2 0 1 3
1630-1659 te) 4 0 1 5
1660-1689 te) 9 10) 0 9
1690-1719 0 5 i) 2 7
1720-1749 t°) 4 i) 0 4
1750-1779 0 10 1 i) 11
1780-1809 0) 2 te) 4 6
1810-1839 i) 8 te) 1 9
1840-1869 2 9 2 3 16
1870-1899 67 16 3 4 90
1900-1929 11 19 3 18 51
1930-1959 37 10 2 6 55
1960— 9 5 3 7 24
No date 25 1 20 27 73
CONTINENTS COMBINED
MOLLUSCS BIRDS MAMMALS OTHER TOTAL TOTAL
40 11 24 46 121 484
te) 0) te) 0 t) 3
0) te) 0 te) 0 5
0 t) te) te) te) 9
i?) te) 0 te) i) 7
te) 0 0 te) 1) 4
0 0 te) (0) 0 11
te) 0 1 (0) 1 7
t°) 1 2 te) 3 12
0 1 1 1 3 19
0 it 6 1 8 98
6 4 3 iu 20 71
25 2 7 15 49 104
4 2 2 12 20 44
5 0 2 10 17 90
Note: these summaries do not take into account 4 species (2 birds, 1 mammal and 1 ‘other’) which are not assignable to either island or continent.
1600. These data should be interpreted cautiously. In only
a few cases are the extinction dates reasonably certain;
more often they are approximate to within one or two
decades. In other cases, they are simply the date when the
species was last recorded, and it is unknown how accurately
they reflect the actual date of extinction (assuming the
species is truly extinct). The uncertainties are most marked
for species in areas which have only been occasionally
surveyed (e.g. land snails on many tropical islands), and
create difficulty in interpreting trends in extinction rates.
Of the individual taxa presented, island birds are the best
documented group. There is no consistent trend over the
full 400 years; peaks occur in the mid-17th and mid-18th
centuries, and there is a clearer increase for the early 19th
century until the 1930s. The apparent fluctuations for the
first 200 years may represent real effects from introduced
species, hunting and habitat modification associated with
increasing levels of human settlement. Continental bird
extinctions and the entire mammal data set are numerically
smaller and thus harder to interpret. Of the 14 dated
mammalian extinctions on islands, 13 have taken place
since 1840. Most of the 20 undated mammalian extinctions
(chiefly Caribbean rodents and insectivores) are believed to
have taken place before the middle of the 19th century,
showing little indication of a marked overall trend.
Information on mollusc extinctions was not available prior
to the mid-19th century, and although high numbers of
extinctions are documented for island molluscs in two 30-
year periods, uncertainty in the dates again confuses
interpretation.
Two trends are apparent in the time-series data for all taxa:
first, that documented island extinctions began almost two
centuries earlier than continental extinctions; second, that
both island and continental extinctions have increased
rapidly from early or mid-19th century to the mid-20th
century. This increase has been more pronounced for
continental species, although the island extinctions exceed
continental ones numerically in all periods. The late 19th
century for islands has the highest rate of all periods,
reflecting a high contribution for mollusc extinctions on
islands during this period.
The apparent decline in rate for both continental and islands
for 1960-1989 is probably attributable to two causes; one is
the expected time-lag in recording extinctions from 1960
onwards. As noted above, extinction is normally only
attributed when a species has not been recorded over a
significant time span. For some purposes, such as the
designation of ‘Extinct’ under the Convention on Trade in
Endangered Species of Wild Fauna and Flora (CITES), this
time period is arbitrarily taken as 50 years. By this
criterion, therefore, no species would be accepted as having
become extinct since 1960 as 50 years would not have
elapsed since its last being recorded. A more realistic and
flexible approach has been adopted here, on the grounds
that some species recorded since 1960 are regarded with a
high degree of certainty to have become extinct, while
conversely many species not observed by specialists in the
wild for over 50 years are almost certainly still extant.
Nevertheless, the general principle holds that the longer a
species has not been recorded the more likely it is to be
regarded as extinct, and vice-versa. A significant number of
species are therefore likely to have become extinct recently
without being recorded as such.
A second, more positive contributory factor to explain the
apparent recent decline in extinction rates is the great
increase in conservation action over the past 30 years.
During this time, attention has focused largely on saving
well-known species under imminent threat of extinction;
most efforts to preserve these species havesucceeded, at
least in the short or medium term. Several projects have
taken the last wild individuals into captivity to build up
populations until environmental conditions and populations
are suitable for re-introduction to the wild (Tables 16.7 and
16.8). Thus well-documented species most vulnerable to
Species Extinction
Figure 16.4 Time series of animal extinctions on islands and continents: selected taxa
Molluscs
60 ;—
40 |;—
20 ;—
no date
Bey ps]
g 1600-1629 1660-1689 1720-1749 1780-1809 1840-1869 1900-1929 1960-
= 1630-1659 1690-1719 1750-1779 1810-1839 1870-1899 1930-1959
te}
iS
=)
zZ .
oo | Birds
=
| no date j 3
1600-1629 1660-1689 1720-1749 1780-1809 1840-1869 1900-1929 1960-
1630-1659 1690-1719 1750-1779 1810-1839 1870-1899 1930-1959
20 Mammals
no
date J
1600-1629 1660-1689 1720-1749 1780-1809 1840-1869 1900-1929 1960-
1630-1659 1690-1719 1750-1779 1810-1839 1870-1899 1930-1959
Pad Islands By continents Year period
201
1. Biological Diversity
Figure 16.5 Time series of animal extinctions on islands and continents: all taxa
100
80 |-
i= L
Ss
z
40 |-
20 |-
no date L
o mam ia i a | x i.
1600-1629 1660-1689 1720-1749 1780-1809 1840-1869 1900-1929
1630-1659 1690-1719 1750-1779 1810-1839 1870-1899 eos
@ islands oy continents Year period
extinction during the past 30 years have often not become
so, as a result of direct manipulative intervention. As noted
above, it seems probable that significant numbers of
undocumented continental species will have become extinct
during this time.
CURRENT AND FUTURE EXTINCTION RATES
Habitat destruction, modification, and fragmentation are
widely recognised as the most serious current threats to
biological diversity, and the primary cause of recent
extinctions. Estimates for present and projected global
extinction rates have not been based on observed or
recorded species extinctions, but rather on extrapolations
from estimates of habitat loss coupled with assumptions
derived from biogeography, relating numbers of species to
area of habitat. A range of estimates are given in
Table 16.3
In practice, most predictions of global extinction rates have
been based on estimates of species richness in tropical
forests, combined with estimates of actual and projected
deforestation rates. Equating global species extinction with
tropical forest species extinction has been justified by the
recognition that the vast majority of terrestrial species occur
in tropical moist forests.
The extrapolations from estimates of habitat loss are
coupled with biogeographic assumptions using the species-
area (Arrhenius) relation (logS = c + zlogA) where S =
number of species, A = area and c and z are constants (see
Chapter 5). Values for z used are between 0.15 and 0.40.
202
The most widely quoted generalisation is that a ten-fold
reduction in area (i.e. loss of 90% of habitat) results in the
loss of half the species present (30% with z = 0.15; 60%
with z = 0.40).
Recent estimates based on these assumptions include those
of Ehrlich and Wilson (1991) and Reid and Miller (1989).
The former, on the basis of a 1.8% loss of rain forest per
year, and using ‘conservative’ estimates from biogeographic
theory (i.e. low z values), estimate a loss of 2-3% of rain
forest species per decade. Reid and Miller, using z values
of 0.15-0.40 and the assumption that forest loss is 1-2 times
that projected by FAO for the period 1980-85, derive a
similar figure of 2-5% loss per decade. This translates into
a loss of some 5-15% by the year 2020, assuming rates of
forest loss continue to increase.
Reid (1992) has refined the analysis somewhat, applying
figures for forest area and rates of loss separately to Latin
America, Africa and Asia, and accounting for observed
differences in species diversity between the three regions.
Using z values of 0.15-0.35 he concludes that global loss of
closed-forest species will be of the order of 1-5% per
decade, or 2-8% in total between 1990 and 2015. Reid
stresses (and this applies to other estimates of species loss)
that this is the number of species ‘committed’ to eventual
extinction as a result of forest loss, not the number which
will actually become extinct during that time - in many
cases, there will be a delay between reduction in area of
habitat and the extinction of species dependent on that
habitat, especially for longer-living species.
METHOD OF ESTIMATION
Table 16.3 Estimated rates of extinction
ESTIMATE % GLOBAL
LOSS PER
DECADE
One million species between 1975 and 4
2000
15-20% of species between 1980 and
2000
12% of plant species in neotropics. -
15% of bird species in Amazon basin
2000 plant species per year in tropics 8
and subtropics
25% of species between 1985 and 9
2015
At least 7% of plant species 7
0.2-0.3% per year 2-3
5-15% forest species by 2020 2-5
2-8% loss between 1990 and 2015 1-5
Extrapolation of past
exponentially increasing trend
Estimated species-area curve;
forest loss based on Global
2000 projections
Species-area curve (z=0.25)
Loss of half the species in area
likely to be deforested by 2015
As above
Half of species lost over next
decade in 10 ‘hot spots’
covering 3.5% of forest area
Half of rain forest species
assumed lost in tropical rain
forests to be local endemics
and becoming extinct with
forest loss
Species-area curve (0.15 < z
<0.35); forest loss assumed
twice rate projected by FAO for
1980-85
Species-area curve (0.15 <z
< 0.35); range includes current
Species Extinction
REFERENCE
Myers (1979)
Lovejoy (1980)
Simberloff (1986)
Raven (1987)
Raven (1988a.b)
Myers (1988)
Wilson (1988,
1989)
Reid and Miller
(1989)
Reid (1992)
rate of forest loss and 50%
increase
Source: Reid, W.V. 1992. How many species will there be? In: Whitmore, T.C. and Sayer, J.A. (Eds), Tropical Deforestation and Species
Extinction, Chapman Hall, London, UK.
Notes: See original source for additional notes referring to this table and reference citations.
Estimates such as these are often combined with estimates
of species numbers in tropical rain forests to provide figures
for numbers of species disappearing daily, yearly or each
decade. Figures of 100,000 species lost per year (based on
estimates of 20 million tropical forest species) are
frequently quoted. The vast majority of the hypothesised
extinctions would occur among undescribed arthropods
because these comprise the majority of the total number of
species estimated to occur in tropical forest.
Earlier estimates, some based on similar biogeographic
assumptions and others using different models, gave even
higher projected rates of extinction, with figures of 20-50%
species loss by the end of the century (Myers, 1979;
Ehrlich and Ehrlich, 1981). In the light of the more recent
estimates based on increased sophistication of the model,
these earlier predictions now look exaggerated.
Problems with the model
Both the theoretical assumptions and the figures used in
deriving estimates from the species-area model are open to
question.
The principal assumption underlying the model is that
species richness and habitat destruction within tropical
203
forests are distributed evenly. This is not the case, as
richness is known to vary considerably between different
areas of tropical moist forest at all scales of comparison
(see Chapter 4). Many ecologists and taxonomists would
agree that, given the inadequate data available on the
poorly-known groups which make up most of the world’s
total complement of species, no realistic assessment can be
made of the extent to which reduction of an area of forest
habitat will affect the species present.
Areas also differ greatly in the number of species confined
to them (i.e. endemics). Self-evidently, the complete
destruction of even a small area with a large number of
endemics will contribute more to global extinction than the
destruction of the same-sized area with few or no local
endemics, even if the latter is richer in species. Thus, if
habitat destruction preferentially takes place in areas with
large numbers of endemics it will lead to extinction rates
higher than those estimated from mean species-area
relationships, while if it is concentrated in areas with few
endemics, the reverse will be the case.
Figures for rates of habitat destruction are also open to
question (see Chapter 20). Calculations tend to take figures
for forest conversion as equivalent to forest loss, that is
complete destruction of forest and replacement by habitats
1. Biological Diversity
in which none of the original biota can survive. In reality,
forest conversion covers a range of conditions, from
selective logging which may have relatively little impact on
species composition, through small-scale patch-work
clearing for agriculture, to clear-felling of extensive areas.
Forest conversion thus covers a range of degrees of
degradation, with only the most extreme resulting in
complete elimination of all species from a particular area.
This will tend therefore to reduce the estimates for
extinction rates. In addition, projections of extinction rates
are based on an assumption that deforestation rates will
remain constant. This is evidently not the case. It is widely
agreed that rates of forest conversion are increasing, and
will continue to increase until easily accessible areas which
are not legally protected have been cleared, following which
they will decrease.
Furthermore, the estimate from a straightforward global
species-area curve does not take into account the presumed
‘residual’ extinctions which will occur through remaining
forest becoming fragmented: on the basis of island
biogeographic theory it is argued that these fragments will
suffer elevated rates of extinction through stochastic
processes. Already many species may be committed to
extinction in that without direct human intervention, their
residual numbers are non-viable. The list of threatened
species in Table 17.1 show 140 species of mammals as
endangered and likely to become extinct in the near future
unless the threat to their survival is alleviated: this is more
than twice the total number of mammals that has gone
extinct over the last four hundred years. Instead of
concentrating on extinctions, it is important to monitor the
status and threats to a wide array of species if global trends
of species diversity are to be assessed.
Finally, estimates of extinction rates do not - and cannot -
take into account the impact of unpredictable large-scale
changes in environmental conditions, such as global climate
change, which is likely to have a profound influence upon
species survival.
Conclusion
There are many unsatisfactory assumptions underlying
current estimates of global extinction rates, and the
resulting numerical values are fraught with imprecision.
Alternative models, possibly based on a_ greater
understanding of the ecological or life history traits
correlated to extinction proneness, would be highly
instructive in either confirming current estimates or refining
them by avoiding some of the major short-comings in the
species - area method. However in the absence of such
alternatives, conclusions from the different studies using the
current model must be examined, even if the methodology
is known to be flawed. In large measure, these agree about
the accelerating rates of species extinctions arising from the
continued loss of tropical forests. The most recent
refinement of the estimates (Reid, 1992) predicts that at
current rates of deforestation, we will commit some 2-8%
of the planet’s species to extinction in the next 25 years.
However, what is equally clear is that quantifying the
precise rate of extinction is of no greater relevance to
conservation practice than is determining a precise figure
for the number of species on earth. Policymakers and the
public may like to assess the magnitude of the extinction
crisis, and thus the priority to be given to the issue, on the
basis of an absolute rate, but investment of time and effort
in refining such predictions contributes little to tackling the
root causes of the problem. Indeed, obsession with an
absolute extinction rate may give an unrealistically
optimistic impression in that no allowance is made for the
genetic impoverishment of the multitude of species brought
to the verge of extinction through the progressive loss of
discrete sub-populations.
Rather than focus on refining extinction rates, we need to
develop the capability to identify areas or localities of high
species endemism and diversity (see Chapter 15), and
ensure that these sites are placed under a system of
conservation management that maintains their ecological
integrity before they are perturbated by logging, mining or
forest clearance. Such proactive conservation practice
could stem the tide of the accelerating species extinction
crisis.
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Conservation Biology: the science of scarcity and diversity.
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Belovsky, G.E. 1987. Extinction models and mammalian persistence.
In: Soulé, M.E. (Ed.), Viable Populations for Conservation.
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Boeckeln, W.J. and Gotelli, N.J. 1984. Island biogeographic theory
and conservation practice: species-area or specious-area
relationships? Biological Conservation 29:63-80.
Bonner, M.L. and Selander, R.K. 1974. Elephant seals: genetic
variation and near extinction. Science 184:908-909.
Brown, J. H. and Kodric-Brown, A. (1977). Turnover rates in insular
biogeography: effect of immigration on extinction. Ecology 58,
445-449.
Ehrlich, P.R. and Ehrlich, A.H. 1981. Extinction: the causes and
consequences of the disappearance of species. Random House,
New York.
Ehrlich, P.R. and Wilson, E.O. 1991. Biodiversity studies: science and
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Erwin, D.H., Valentine, J.W. and Sepkoski, J.J. 1987. A comparative
study of diversification events: the early Palaeozoic versus the
Mesozoic. Evolution 41(6).
Falconer, D.S. 1981. Introduction to Quantitative Genetics, 2nd
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Karr, J.R. 1991. Avian survival rates and the extinction process on
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Klein, B.C. 1989. Effects of forest fragmentation on dung and carrion
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Species Extinction
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Based on text prepared by Martin Jenkins with additions by
WCMC staff
1. Biological Diversity
Table 16.4 Animal species extinct since circa 1600
SPECIES
CORALS ETC. (CNIDARIA)
Order MILLEPORINA
Family Milleporidae
Millepora sp.
MOLLUSCS
Order ARCHAEOGASTROPODA
Family Acmaeidae
Lottia alveus
Order MESOGASTROPODA
Family Hydrobiidae
Bythiospeum pfeifferi
Clappia umbilicata
Ohridohauffenia drimica
Family Pleuroceridae
Elimia clausa
Elim fusiformis
Elimia hartmaniana
Elim impressa
Elima jonesi
Elima laeta
Elima pilsbryi
Elimia pupaeformis
Elima pygmaea
Elima varians
Gyrotoma incisa
Gyrotoma lewisi
Gyrotoma pagoda
Gyrotoma pumila
Gyrotoma pyramidata
Gyrotoma walkeri
Leptoxis clipeata
Leptoxis formanii
Leptoxis ligata
Leptoxis lirata
Leptoxis occultata
Leptoxis showalterii
Leptoxis vittata
Family Pomatiasidae
Tropidophora carinata
Order STYLOMMATOPHORA
Family Endodontidae
Discus guerinianus
Kondoconcha othnius
Libera subcavernula
Libera tumuloides
Mautodonia acuticosta
Mautodonta boraborensis
Mautodonta ceuthma
Mautodonta consimilis
Mautodonta consobrina
Mautodonta maupiensis
Mautodonta parvidens
Mautodonta punctiperforata
Mautodonia saintjohni
Mautodonta subtilis
Mautodonia unilameliata
Mautodonta zebrina
Opanara altiapica
Opanara areaensis
- Opanara bitridentata
+ Opanara caliculata
+ Opanara depasoapicata
+ Opanara duplicidentata
+ Opanara fosbergi
+ Opanara megomphala
« Opanara perahuensis
+ Orangia cooki
+ Orangia maituatensis
+ Orangia sporadica
* Pilula cycloria
Rhysoconcha atanuiensis
Rhysoconcha variumbilicata
Ruatara koarana
Ruatara oparica
Taipidon anceyana
Talpidon marquesana
Talpidon octolamellata
Thaumatodon multilamellatus
Family Bulimulidae
Amphibulima patula
Bulimulus duncanus
Leuchocharis loyaltyensis
Leuchocharis porphyrocheila
ENGLISH NAME
Eelgrass Limpet
Umbilicate Pebblesnail
Closed Elimia
Fusiform Elimia
High—spired Elimia
Constricted Elimia
Hearty Elimia
Ribbed Elimia
Rough-lined Elimia
Pupa Elimia
Pygmy Elimia
Puzzle Elimia
Excised Slitshell
Striate Slitshell
Pagoda Slitshell
Ribbed Slitshell
Pyramid Slitshell
Round Slitshell
Agate Rocksnail
Interrupted Rocksnail
Rotund Rocksnail
Lirate Rocksnail
Bigmouth Rocksnail
Coosa Rocksnail
Striped Rocksnail
206
DISTRIBUTION
Panama
USA
Austria
USA
Yugoslavia
USA
USA
USA
USA
USA
USA
USA
USA
USA
USA
USA
USA
USA
USA
USA
USA
USA
USA
USA
USA
USA
USA
USA
Mauritius
Madeira (Portugal)
Rapa (F. Paynes)
Raratonga (Cook Is)
Raratonga (Cook Is)
Raiatea (F. Polynesia)
Borabora (F. Polynesia)
Raivavae (F. Polynesia)
Raiatea (F. Polynesia)
Huahine (F. Polynesia)
Maupiti (F. Polynesia)
Society Is (F. Polynesia)
Moorea (F. Polynesia)
Borabora (F. Polynesia)
Huahine (F. Polynesia)
Raratonga (Cook Is)
Raratonga (Cook Is)
Rapa (F. Polynesia)
Rapa (F. Polynesia)
Rapa (F. Polynesia)
Rapa (F. Polynesia)
Rapa (F. Ei diet
Rapa (F. Polynesia
Rapa (F. Polynesia)
Rapa (F. Polynesia)
Rapa (F. Se des
Rapa (F. Polynesia)
Rapa (F. Polynesia
Rapa (F. Polynesia
Rapa (F. Polynesia)
Rapa (F. Polynesia
Rapa (F. Polynesia)
Rapa (F. Polynesia)
Hiva Oa (F. Polynesia)
Nuku Hiva (F. Polynesia)
Hiva Oa (F. Polynesia)
Raratonga (Cook Is)
Guadeloupe
Galapagos (Ecuador)
New Caledonia
New Caledonia
LAST
1983
1980s
1924
1924
1924
1924
1924
1924
1881
1870s
1934
1880s
1880s
1880s
1880s
1880s
1880s
1880s
1880s
1880s
1880s
1880s
1880s
1880s
1880s
1934
1934
1934
1934
1934
1934
1934
1934
1934
1934
1934
1934
1934
1934
1934
1934
1880s
1880s
1880s
1880s
late 1800s
1900s
1900s
POSSIBLE
RECORDED CAUSE
fecfo:ferereriesherierherier)
onorm
Species Extinction
Table 16.4 Animal species extinct since circa 1600 (continued)
SPECIES ENGLISH NAME DISTRIBUTION LAST POSSIBLE
RECORDED CAUSE
MOLLUSCS (continued)
Family Charopidae
Helenoconcha leptalea St Helena 1870s
Helenoconcha minutissima St Helena 1870s
Helenoconcha polyodon St Helena 1870s
Helenoconcha pseustes St Helena 1870s
Helenoconcha sexdentata St Helena 1870s
Helenodiscus bilamellata St Helena 1870s
Helenodiscus vernoni St Helena 1870s
Pseudohelenoconcha dianae St Helena 1870s
Pseudohelenoconcha laetissima St Helena 1870s
Pseudohelenoconcha persoluta St Helena 1870s
Pseudohelenoconcha spurca St Helena 1870s
Sinployea canalis Raratonga (Cook Is 1872
Sinployea decorticata Raratonga (Cook Is 1872
Sinployea harveyensis Raratonga (Cook Is 1872
Sinployea otareae Raratonga (Cook Is 1872
Sinployea planospira Raratonga (Cook Is 1872
Sinployea proxima Raratonga (Cook Is 1872
Sinployea rudis Raratonga (Cook Is 1872
Sinployea tenuicostata Raratonga (Cook Is 1872
Sinployea youngi Raratonga (Cook Is 1872
Family Achatinellidae
Achatinell abbreviata Hawaii (USA) 1963 A,C,D
Achatinella buddii Hawaii (USA) early1900s A,C,D
Achatinella caesia Hawaii (USA) early 1900s A,C,D
Achatinella casta Hawaii (USA) A,C,D
Achatinella decora Hawaii (USA early 1900s A,C,D
Achatinella elegans Hawaii (USA 1952 A,C,D
Achatinella juddii Hawaii (USA 1958 A,C,D
Achatinella juncea Hawaii (USA) A,C,D
Achatinella lehuiensis Hawaii (USA) 1922 A,C,D
Achatinella papyracea Hawaii (USA) 1945 A,C,D
Achatinella rosea Hawaii (USA) 1949 A,C,D
Achatinella spaldingi Hawaii (USA) 1938 A,C,D
Achatinella stewarti Hawaii (USA) 1961 A,C,D
Achatinella thaanumi Hawaii (USA) 1900s A,C,D
Achatinell valida Hawaii (USA 1951 A,C,D
Achatinelk vittata Hawaii (USA 1953 A,C,D
x Elasmas jaufreti Rodrigues (Mauritius)
x Elasmas sp. Mauritius
Partulina crassa Hawaii (USA) 1914 Cc?
Partulina montagui Hawaii (USA) 1913 Cc?
Family Partulidae
Partula exigua Moorean Viviparous Tree Snail Moorea (F. Polynesia) 1977 Cc?
Partula filosa Tahiti Viviparous Tree Snail Tahiti (F. Polynesia)
Partula producta Tahiti Viviparous Tree Snail Tahiti (F. Polynesia)
Partula salifana Guam
Samoana abbreviata American Samoa 1940 B
Family Amastridae
Carelia anceophila Hawaii (USA) 1930 B,c,D
Carelia bicolor Hawaii (USA) 1970 B,C,D
Carelia cumingiana Hawaii (USA) 1930 B,C,D
Carelia glossema Hawaii (USA 1930 B,C,D
Carelia kalalauensis Hawaii (USA) 1945/47 B,C,D
Carelia knudseni Hawaii (USA) 1930 B,C,D
Carelia olivacea Hawaii (USA 1930 B,C,D
Carelia paradoxa Hawaii (USA) 1930 B,C,D
Carelia periscelis Hawaii (USA) 1930 B,C,D
Carelia tenebrosa Hawaii (USA) 1930 B,C,D
Carelia turricula Hawaii (USA) 1930 B,C,D
Family Vertiginidae
Campolaemus perexilis St Helena 1870s
Nesopupa turtoni St Helena 1870s
Family Pupillidae
x Gibbulinopss sp. Rodrigues (Mauritius)
Leiostyk abbreviata Madeira (Portugal) 1870s
Leiostyla cassida Madeira (Portugal) 1870s
Leiostyla concinna Madeira (Portugal) 1870s
Leiostyle gibba Madeira (Portugal) 1870s
Leiostyk laevigata Madeira (Portugal) 1870s
Leiostyk lamellosa Madeira eee 1870s
Leiostyf simulator Madeira (Portugal) 1870s
Pupa obliquicostata St Helena 1870s
Family Helixarionidae
Colparion madgei Rodrigues (Mauritius) 1938 B
Ctenoglypta newtoni Mauritius 1871 B
x Ctenophila planorbina Mauritius
Diastole matafaoi American Samoa 1940 2D
x Erepta thiriouxi Mauritius
x Erepta sp. Mauritius
Pachystyla ruforonata Mauritius 1869 B
x Plegma bewsheri Rodrigues (Mauritius)
x Plegma duponti Mauritius
x Plegma sp. Mauritius
Family Ferussaciidae
Cecilioides eulima Madeira (Portugal) 1870s
Family Subulinidae
Chilonopsis blofeldi St Helena 1870s
Chilonopsis exulatus St Helena 1870s
Chilonopsis helena St Helena 1870s
1. Biological Diversity
Table 16.4 Animal species extinct since circa 1600 (continued)
SPECIES ENGLISH NAME DISTRIBUTION LAST POSSIBLE
RECORDED CAUSE
MOLLUSCS (continued)
Chilonopsis melanoides St Helena 1870s
Chilonopsis nonpareil St Helena 1870s
Chilonopsis subplicatus St Helena 1870s
Chilonopsis subtruncatus St Helena 1870s
Chilonopsis turtoni St Helena 1870s
Family Helicidae
Discuk lyelliana Madeira (Portugal) 1870s
Discuk tetrica Madeira (Portuga 1870s
Geomitra delphinuloides Madeira (Portugal 1870s
Lemniscia galeata Madeira (Portugal 1870s
Pseudocampylaea lowei Madeira (Portugal; late 19th C
Family Streptaxidae
Edentulina thomasetti Seychelles 1908
Gibbus lyonetianus Mauritius 1905 B
Gonidomus newtoni Mauritius 1867 B
x Gonospira cirneensis Mauritius
x Gonospira heliodes Mauritius
x Gonospira majusculus Mauritius
Imperturbata violescens? Seychelles
Family Assimineidae
x Omphalotropis plicosa Mauritius 1878 B
x Omphalotropis caldwelli Mauritius
x Omphalotropis dupontiana Mauritius
x Omphalotropis maxima Mauritius
x Omphalotropis muttiirata Mauritius
x Omphalotopis sp. Mauritius
Family Pomatiasidae
x Tropidophora bewsheri Rodrigues (Mauritius)
x Tropidophora bipartita Rodrigues (Mauritius)
x Tropidophora defloraia Réunion
x Tropidophora lienardi Mauritius
x Tropidophora mauritiana Mauritius
Order UNIONOIDA
Family Unionidae
Alasmidonta mecordi Coosa Elktoe USA
Alasmidonta wrightiana Ochlacknee Arc—mussel USA
cesta arcaeformis Sugarspoon USA 1940s B
a arse biemarginata Angled Riffleshell USA 1960s B
‘pioblasma flexuosa Leafshell USA 1940s B
Barrera haysiana Acornshell USA
Epiopiesms lenior Narrow Catspaw USA 1965 B
Farrel lewisi Forkshell USA 1964 B
Eplcbisae. personata Round Combshell USA 1930 B
Epos propinqua Tennessee Riffleshell USA 1930 B
‘pioblasma sampsoni Wabask Riffleshell USA 1950s/60s B
Epioblasma stewardsoni Cumberland Leafshell USA 1930 B
Medionidus meglameriae Tombigbee Moccasinshell USA
CRUSTACEANS
Order AMPHIPODA
Family Crangonyctidae
Stygobromus hayi Hay’s Spring Scud USA 1957
Stygobromus lucifugus Rubious Cave Amphipod USA
Order DECAPODA
Family Astacidae
Pacifastacus nigrescens Sooty Crayfish USA 1860s
Family Atyidae
Syncaris pasadenas Pasadena Freshwater Shrimp USA 1933
INSECTS
Order EPHEMEROPTERA
Family Siphlonuridae
Acanthometropus pecatonia Pecatonica River Mayfly USA 1927
Family Ephemeridae
Pantagenia robusta Robust Burrowing Mayfly USA
Order ORTHOPTERA
Family Tettigoniidae
Neduba extincta Antioch Dunes Shieldback Katydid USA 1937
Order PHASMATOPTERA
Family Phasmatidae
Dryococelus australis Lord Howe Island Stick—insect Lord Howe | (Australia) 1969
Order DERMAPTERA
Family Labiduridae
* Labidura herculeana St Helena Earwig St Helena 1967
Order PLECOPTERA
Family Chloroperiidae
Alloperla roberti Robert's Stonefly USA
208
Table 16.4 Animal species extinct since circa 1600 (continued)
SPECIES
INSECTS (continued)
Order HOMOPTERA
Family Pseudococcidae
Clavicoccus erinaceus
Phyllococcus oahuensis
Order COLEOPTERA
Family Cerambycidae
Xyloteles costatus
Family Curculionidae
Dryophthorus distinguendus
Dryotribus mimeticus
Hadramphus tuberculatus
Macrancylus linearis
Oedemasylus laysanensis
Pentarthrum blackburnii
Rhyncogonus bryani
Family Carabidae
* Aplothorax burchelli
* Mecodema punctellum
Order DIPTERA
Family Tabanidae
Stonemyia volutina
Family Dolichopodidae
Campsicnemus mirabilis
Family Drosophilidae
Drosophila lanaiensis
Order TRICHOPTERA
Family Rhyacophilidae
Rhyacophila amabilis
Family Hydropsychidae
Hydropsyche tobiasi
Family Leptoceridae
Triaenodes phalacris
Triaenodes tridonata
Order LEPIDOPTERA
Family Zygaenidae
Levuana iridescens
Family Lycaenidae
Glaucopsyche xerces
Family Libytheidae
Libythea cinyras
Family Nymphalidae
Euthalia malapana
Family Pyralidae
Genophantis leahi
Hedylepta asaphombra
Heaylepta coninuatalis
Hedylepta epicentra
+ Hedylepta euryprora
+ Heaylepta fullawayi
Hedylepta laysanensis
+ Hedylepta meyricki
+ Hedylepta musicola
Hedylepta telegrapha
Ocobia sp.
Family Geometridae
Scotorhythra nesiotes
Scotorhythra megalophylla
Scotorhythra paratactis
Tritocleis microphylla
Family Sphingidae
Mandura blackburni
Family Noctuidae
Agrotis crinigera
Agrotis fasciata
Agrotis kerri
Agrotis laysanensis
Agrotis pi ila
Agrotis procellaris
Helicoverpa confusa
Helicoverpa minuta
5 dca laysanensis
ar es newelli
+ fe plagiota
+ Hypena senicula
Peridroma porphyrea
Order HYMENOPTERA
Family Colletidae
Nesoprosopis angustula
Nesoprosopis blackburni
Nesoprosopis connectens
ENGLISH NAME
Pitt lsland Longhorn Borer
Volutine Stoneyian Tabanid Fly
Castle Lake Caddis —fly
Tobias’ Caddis —fly
Athens Caddis—fly
Three—tooth Caddis —fly
Levuana Moth
Xerces Blue
Poco Noctuid Moth
Midway Noctuid Moth
Minute Noctuid Moth
Laysan Dropseed Noctuid Moth
Lanai Yellow—faced Bee
Blackburn’s Yellow—faced Bee
Connected Yellow—faced Bee
209
DISTRIBUTION
Hawaii (USA)
Hawaii (USA)
Chatham | (NZ)
Hawaii (USA)
Hawaii (USA)
New Zealand
Hawaii (USA
Hawaii (USA)
Hawaii (USA)
Hawaii (USA)
St Helena
Stephens | (NZ)
USA
Hawaii (USA)
Hawaii (USA)
USA
Germany
USA
USA
Fiji
USA
Mauritius
Taiwan
Hawaii (USA
Hawaii (USA)
Hawaii (USA)
Hawaii (USA\
Hawaii (USA)
Hawaii (USA\
Hawaii (USA)
Hawaii (USA)
Hawaii (USA)
Hawaii (USA)
Hawaii (USA)
Hawaii (USA)
Hawaii (USA)
Hawaii (USA
Hawaii (USA
Hawaii (USA'
Hawaii (USA'
Hawaii (USA)
Hawaii (USA'
Hawaii (USA
Hawaii (USA
Hawaii (USA
Hawaii (USA
Hawaii (USA)
Hawaii (USA)
Hawaii SA
Hawaii (USA)
Species Extinction
LAST POSSIBLE
RECORDED CAUSE
1930s B,C
1910 c
1967?
BiG
1920s
1929 E
early 1940s
1865
early 1900s
1970s
1958
early 1900s
mm mm
1911
early 1900s
early 1900s
early 1900s
1890s
1960s
1926 E
1923
1911
pre—1900
post—1927
pre—1911
1911
1. Biological Diversity
Table 16.4 Animal species extinct since circa 1600 (continued)
SPECIES
FAISHES
Order PETROMYZONTIFORMES
Family Petromyzontidae
Lampeta minima
Order CYPRINIFORMES
Family Cyprinidae
Evarra bustamantei
Evarra eigenmanni
Evarra tahuacensis
Gila crassicauda
Lepidomeda attivelis
Notropis amecae
Notropis aulidion
Notropis orca
Pogonichthys ciscoides
Rhinichthys deaconi
Stypodon signifer
Family Catostomidae
Chasmistes muriei
Lagochila lacera
Order SALMONIFORMES
Family Retropinnidae
* Prototroctes oxyrhynchus
Family Salmonidae
Coregonus alpenae
Coregonus johannae
Salvelinus agassizi
Order CYPRINODONTIFORMES
Family Fundulidae
Fundulus albolineatus
Family Poeciliidae
Gambusia amistadenss
* Gambusia georgei
* Priapella bonita
Family Goodeidae
Characodon garmani
Empetrichthys merriami
Family Cyprinodontidae
Cyprinodon latifasciatus
Cyprinodon sp.
Cyprinodon sp.
Cyprinodon sp.
Order SCORPAENIFORMES
Family Cottidae
Cottus echinatus
AMPHIBIANS
Order ANURA
Family Discoglossidae
Discoglossus nigriventer
Rana fisheri
REPTILES
Order TESTUDINES
Family Testudinidae
Cylindraspis borbonica
Cylindraspis indica
Cylindraspis inept
Cylindraspis peltastes
Cylindraspis triserrata
Cylindraspis vosmaeri
Order SAURIA
Family Gekkonidae
Hoplodactylus delcourti
Phelsuma edwardnewtoni
Phelsuma gigas
Family Iguanidae
Leiocephalus eremitus
Leiocephalus herminieri
Family Teiidae
Ameiva cineracea
* Ameiva major
Family Anguidae
Celestus occiduus
Family Scincidae
# Leiolopisma mauritiana
Macroscincus coctei
* Tiliqua adelaidensss
ENGLISH NAME
Miller Lake Lamprey
Thicktail Chub
Pahranagat Spinedace
Ameca Shiner
Durango Shiner
Phantom Shiner
Clear Lake Splittail
Las Vegas Dace
Stumptooth Minnow
Snake River Sucker
Harelip Sucker
New Zealand Grayling
Longjaw Cisco
Deepwater Cisco
Silver Trout
Whiteline Topminnow
Amistad Gambusia
San Marcos Gambusia
Guayacon Ojiazul
Parras Characodon
Ash Meadows Killifish
Perrito de Parras
Monkey Spring Pupfish
Utah Lake Sculpin
Israel Painted Frog
Relict Leopard Frog
Newton's Day Gecko
Giant Day Gecko
Martinique Giant Ameiva
Jamaican Giant Galliwasp
Cape Verde Giant Skink
Adelaide Pigmy Bluetongue
DISTRIBUTION
USA
Mexico, USA
USA
USA
Mexico
USA
USA
New Zealand
USA, Canada
USA, Canada
USA
USA
USA
USA
Mexico
Mexico
USA
Mexico
USA
Mexico
Mexico
USA
Israel
USA
Réunion
Réunion
Mauritius
Rodrigues (Mauritius)
Mauritius
Rodrigues (Mauritius)
New Zealand (?)
Rodrigues es
Rodrigues (Mauritius
Navassa | (USA)
Martinique
Guadeloupe
Martinique
Jamaica
Mauritius
Cape Verde
Australia
LAST POSSIBLE
RECORDED CAUSE
1953 E
1970 B
1970 B
1970 B
1957 B,C/D
1940 c/D
1970 c/D
1965 c/D
1975 B,C/D
1970 B,c/D
1955 B
1930 B
1928 B
1910 G
1920s B,D,H
1978 A,C
1955 A,C/D
1930 A,C/D
1900 B,c/D
1973 B
1983 B,C/D
1900 2B
1953 B,C/D
1930 B
1971 B,C/D
1928 B,C/D
1940 B
1960 B
1800
1800 A
early 18th C A,C/D
1800 A,B,C/D
early 18thC A,C/D
1800 A,C/D
mid 19th C?
1917 Cc
endi9thC C
1900 c
1830s —
early 20thC —
Cc?
1840 Cc?
1600 c
early 20thC A?
1959 B,C
Species Extinction
Table 16.4 Animal species extinct since circa 1600 (continued)
SPECIES ENGLISH NAME DISTRIBUTION LAST POSSIBLE
RECORDED CAUSE
REPTILES (continued)
Order SERPENTES
Family Boidae
* Bolyeria muttocarinata Round | (Mauritius) 1975
Family Typhlopidae
Typhlops cariei Mauritius 17th C Cc
Family Colubridae
* Alsophis ater Jamaican Tree Snake Jamaica 1950 A,c
Alsophis sancticrucis St Croix Racer Virgin Is (US) 20th C A.C
* Liophis cursor Martinique Racer Martinique 1963 Cc
* Liophis perfuscus Barbados Racer Barbados mid 20th C? C
BIRDS:
Order STRUTHIONIFORMES
Family Dromaiidae
Dromaius diemenianus Kangaroo Island Emu Kangaroo | (Australia) 1803 B
Family Aepyornithidae
Aepyornis maximus Great Elephantbird Madagascar 1650 A.B
Family Anomalopterygidae
Dinornis torosus Brawny Great Moa New Zealand 1670 A.B
Eurapteryx gravis Burly Lesser Moa New Zealand 1640 A,B
Megalaperyx didinus South Island Tokoweka New Zealand 1785 A,B
Order GALLIFORMES
Family Phasianidae
Coturnix novaezelandiae New Zealand Quail New Zealand 1875 F
Ophrysia superciliosa Himalayan Mountain Quail India 1868 A
Order ANSERIFORMES
Family Anatidae
Alopochen mauritianus Mauritian Shelduck Mauritius 1698 =
Anas theodori Mauritian Duck Mauritius, 7Réunion 1696 =
Camptorhynchus labradorius Labrador Duck Canada, USA 1878 A,B
Cygnus sumnerensis Chatham Island Swan Chatham | (NZ) 1590-1690 —
gus australis Auckland Island Merganser New Zealand 1905 A,B,C
* Rhodonessa caryophyllacea Pink—headed Duck India, Nepal 1935 A
Sheldgoose sp. Réunion 1674 =
Order CORACIIFORMES
Family Alcedinidae
Halcyon miyakoensis Ryukyu Kingfisher Nansei—shoto (Japan) 1841 =
Order CUCULIFORMES
Family Cuculidae
* Coua delalandei Snail—eating Coua Madagascar 1930 A,B,C/D
Order PSITTACIFORMES
Family Psittacidae
Anodorhynchus glaucus Glaucous Macaw Brazil, Uruguay 1955
Ara tricolor Cuban Red Macaw Cuba 1885 AE
Charmosyna diadema New Caledonia Lorikeet New Caledonia 1860 B
Conuropss carolinensis Carolina Parakeet USA 1914 E
Cyanoramphus ulietanus Raiatea Parakeet Raiatea (F. Polynesia) 1773 =
Cyanoramphus zealandicus Black—fronted Parakeet Tahiti (F. Polynesia) 1844 B
Lophopsittacus’ bensoni Mauritius Grey Parrot Mauritius 1765 c/D
Lophopsittacus mauritianus Mauritius Parrot Mauritius 1675 A,C
Mascarinus mascarinus Mascarene Parrot Réunion 1775 (1834 B
in captivity)
'Necropsittacus 'rodericanus Rodrigues Parrot Rodrigues (Mauritius) 1761 A,C/D
Nestor productus Norfolk Island Kaka Phillip | (Australia) 1851 AE
Psittacula exsul Rodrigues Ring—necked Parakeet Rodrigues (Mauritius) 1876 B
Psittacula wardi Seychelles Alexandrine Parrot Seychelles 1870 A.B
Order TROCHILIFORMES
Family Trochilidae
Chlorostilbon bracei New Providence Hummingbird Bahamas 1877
Family Caprimulgidae
* Siphonorhis americanus Jamaica Least Pauraque Jamaica 1859 Cc
Order STRIGIFORMES
Family Strigidae
Athene blewitti Forest Owet India 1914
‘Athene’ murivora Rodrigues Little OW Rodrigues (Mauritius) 1726 B
?Sauzieri sp. Mauritian OW Mauritius
* Sceloglaux albifacies Laughing OM New Zealand 1914 B,C
‘Scops’ commersoni Mauritian OW Mauritius 1836
Family Aegothelidae
* Aegotheles savesi New Caledonia Owet—frogmouth New Caledonia 1880 =
Order COLUMBIFORMES
Family Raphidae
‘Ornithaptera’ solitaria Réunion Solitare Réunion 1710-1715 A
Pezophaps solitarius Rodrigues Solitare Rodrigues (Mauritius) 1765 A
Raphus cucullatus Dodo Mauritius 1665 A,C,D
1. Biological Diversity
Table 16.4 Animal species extinct since circa 1600 (continued)
SPECIES
BIRDS (continued)
Family Columbidae
Alextroenas nitidissima
‘Alextroenas' rodericana
Columba jouyi
Columba versicolor
Ectopistes migratorius
* Microgoura meeki
* Ptilinopus mercierii
Order GRUIFORMES
Family Rallidae
Aphanapteryx bonasia
Aphanapteryx leguati
Atlantisia elpenor
Fulica newtoni
Gallinula nesiotis
Gallinula pacifica
Gallirallus pacificus
Nesoclopeus woodfordi
Porphyrio albus
Porzana monasa
Porzana palmeri
Porzana sandwichensis
Rallus dieffenbachi
Rallus modestus
Rallus wakensis
* Tricholimnas lafresnayanus
Order CICONIIFORMES
Family Scolopacidae
Prosobonia leucoptera
Family Charadriidae
Haematopus meadewaldoi
Vanellus macropterus
Family Laridae
Alca impennis
Family Falconidae
Falco sp.
Polyborus lutosus
Family Podicipedidae
Podiceps andinus
Podilymbus gigas
Tachybaptus rufolarvatus
Family Phalacrocoracidae
Phalacrocorax perspicillatus
Family Ardeidae
lxobrychus novaezelandia
Nycticorax mauritianus
Nycticorax megacephalus
Nycticorax sp.
Family Threskiornithidae
Borbonibis latipes
Family Ciconiidae
Ciconia sp.
Family Procellariidae
* Oceanodroma macrodactyla
Pterodroma sp.
Order PASSERIFORMES
Family Acanthisittidae
Xenicus longipes
Xenicus lyalli
Family Pycnonotidae
Hypsipetes sp.
Family Muscicapidae
Acrocephalus familiaris
Eutrichomyas rowleyi
Myiagra freycineti
* Turnagra capensis
Turdus ravidus
Zoothera terrestris
Babbler sp.
Family Dicaeidae
Dicaeum quadricolor
Family Zosteropidae
Zosterops strenua
Family Meliphagidae
Chaetoptila angustipluma
Moho apicalis
* Moho nobilis
ENGLISH NAME
Pigeon Hollandais
Rodrigues Pigeon
Ryukyu Wood Pigeon
Bonin Wood Pigeon
Passenger Pigeon
Solomon Island Crowned—pigeon
Marquesas Fruit—dove
Red Rail
Rodrigues Rail
Ascension Flightless Crake
Mascarene Coot
Tristan Moorhen
Samoan Woodhen
Tahiti Rail
Woodford's Rail
Lord Howe Purple Gallinule
Kosrae Crake
Laysan Rail
Hawaiian Rail
Chatham Island Banded Rail
Chatham Island Rail
Wake Island Rail
New Caledonia Rail
Tahitian Sandpiper
Canarian Black Oystercatcher
Javanese Wattled Lapwing
Great Auk
Guadalupe Caracara
Colombian Grebe
Atitlan Grebe
Lake Alaotra Grebe
Spectacled Cormorant
New Zealand Little Bittern
Mauritius Night—heron
Rodrigues Night—heron
Reunion Flightless Ibis
Guadalupe Storm—petrel
Bush Wren
Stephens Island Wren
Laysan Millerbird
Caerulean Paradise—flycatch
Guam Broadbill
Piopio
Grand Cayman Thrush
Kittlitz’s Thrush
Four —coloured Flowerpecker
Lord Howe White—eye
Kioea
Oahu Oo
Hawaii Oo
i)
=
tN
DISTRIBUTION
Mauritius
Rodrigues (Mauritius)
Nansei—shoto (Japan)
Ogasawara—shoto (Japan)
USA
Choiseul (Solomon Is)
Marquesas Is (F. Polynesia)
Mauritius
Rodrigues (Mauritius)
Ascension | (UK)
Mauritius, Reunion
Tristan da Cunha (UK)
Savaii (Western Samoa)
French Polynesia
Bougainville (Papua New Guinea)
Lord Howe | (Australia)
Federated States of Micronesia
Hawaii (USA)
Hawaii (USA)
Chatham | (NZ)
Chatham | (NZ)
Wake | (USA)
New Caledonia
Tahiti, Moorea (F. Polynesia)
Canary Is (Spain)
Java (Indonesia)
Canada, Iceland, Faeroes
UK, ‘USSR’, Greenland
Réunion
Guadalupe (Mexico)
Colombia
Guatemala
Madagascar
Bering Straits (‘USSR’)
New Zealand
Mauritius
Rodrigues (Mauritius)
Réunion
Réunion
Réunion
Guadalupe (Mexico)
Rodrigues (Mauritius)
New Zealand
Stephens | (NZ)
Rodrigues (Mauritius)
Hawaii (USA)
Sangihe (Indonesia)
Guam
New Zealand
Cayman Is
Ogasawara—shoto (Japan)
Rodrigues (Mauritius)
Cebu (Philippines)
Lord Howe | (Australia)
Hawaii (USA)
Hawaii (USA)
Hawaii (USA)
LAST
POSSIBLE
RECORDED CAUSE
1835
1726
1936
1889
1914
1904
1922
1700
1761
1656
1693
1875-1900
1908-1926
1773-4
1936
1834
1827
1944
1898
1840
1900
1945
1904
1773
1913
1940
1844
1674
1900
1977
ie]
| >omng00>|
re)
A,D,E
1980—1986/7A,D
1852
1900
by 1700
1761
by 1700
1773
1674
1912-1922
1726
1972
1874
1600s?
1912-1923
1978
1983
1955
1938
1928
1600s?
1906
1928
1860
1837
1934
Table 16.4 Animal species extinct since circa 1600 (continued)
SPECIES
BIRDS (continued)
Ciridops anna
Drepanis funerea
Drepanis pacifica
* Hemignathus obscurus
Hemignathus sagittirostris
* Paroreomyza flammea
Psittrostra kona
Rhodacanthis flaviceps
Rhodacanthis palmeri
Family Icteridae
Quiscalus palustris
Family Ploceidae
Foucta sp.
Family Fringillidae
Chaunoproctus ferreorostris
Spiza townsendi
Family Sturnidae
Aplonis corvina
Aplonis fusca
Aplonis mavornata
* Aplonis pelzelni
Tegilupus varius
Necrospar rodericanus
Family Callaeidae
Heteralocha acutirostris
MAMMALS
Order MARSUPIALIA
Family Macropodidae
* Caloprymnus campestris
+ Lagorchestes asomatus
Lagorchestes leporides
Macropus greyi
Onychogalea lunata
Potorous platyops
Family Peramelidae
Chaeropus ecaudatus
Perameles eremiana
Family Thylacomyidae
Macrotis leucura
Family Thylacinidae
Thylacinus cynocephalus
Order CHIROPTERA
Family Pteropodidae
Acerodon lucifer
Dobsonia chapmani
Pteropus pilosus
Pteropus subniger
Pteropus tokudae
Family Molossidae
Mystacina robusta
Order INSECTIVORA
Family Nesophontidae
# Nesophontes hypomicrus
# Nesophontes micrus
# Nesophontes paramicrus
# Nesophontes zamicrus
# Nesophontes sp.
Order LAGOMORPHA
Family Ochotonidae
Prolagus sardus
Family Leporidae
* Sylviagus insonus
Order RODENTIA
Family Arvicolidae
Pitymys bavaricus
Family Capromyidae
# Capromys sp.
# Geocapromys colombianus
Geocapromys thoractus
# Geocapromys sp.
# Isolobodon portoricensis
# Plagiodontia velozi
Family Cricetidae
Megalomys desmarestii
Megalomys luciae
Megaloryzomys curioi
Megaloryzomys sp.
Nesoryzomys darwini
Nesoryzomys sp.
ENGLISH NAME
Ula—ai—hawane
Black Mamo
Hawaii Mamo
Akialoa
Greater Amakihi
Kakawihie or Molokai Creeper
Kona Grosbeak
Lesser Koa—finch
Greater Koa—finch
Slender —billed Grackle
Reunion Fody
Bonin Grosbeak
Townsend's Finch
Kosrae Mountain Starling
Norfolk Island Starling
Mysterious Starling
Pohnpei Mountain Starling
Réunion Starli
Rodrigues Starling
Huia
Desert Rat—kangaroo
Central Hare—wallaby
Eastern Hare—wallaby
Toolache Wallaby
Crescent Nailtail Wallaby
Broad—faced Potoroo
Pig—footed Bandicoot
Desert Bandicoot
Lesser Bilby
Thylacine
Panay Giant Fruit Bat
Chapman's Bare—backed Flying Fox
Palau Flying Fox
Lesser Mascarene Flying Fox
Guam Flying Fox
New Zealand Lesser Short-—tailed Bat
Atalaye Nesophontes
Western Cuban Nesophontes
St Michel Nesophontes
Haitian Nesophontes
Sardinian Pika
Omilteme Cottontail
Bavarian Pine Vole
Martinique Rice Rat
St Lucia Rice Rat
Santa Cruz Rice Rat
DISTRIBUTION
Hawaii (USA)
Hawaii (USA
Hawaii (USA
Hawaii (USA
Hawaii (USA\
Hawaii (USA‘
Hawaii (USA)
Hawaii (USA\
Hawaii (USA)
Mexico
Réunion
Ogasawara—shoto (Japan)
USA
Kosrae (Fed. States Micronesia)
Norfolk | (Australia)
Cook Is
Pohnpei (Fed. States Micronesia)
Réunion
Rodrigues (Mauritius)
New Zealand
Australia
Australia
Australia
Australia
Australia
Australia
Australia
Australia
Australia
Tasmania (Australia)
Philippines
Philippines
Palau
Mauritius, Réunion
Guam
New Zealand
Haiti, Dominican Republic
Cuba
Haiti, Dominican Republic
Haiti, Dominican Republic
Cayman Is
Corsica (France), Sardinia (Italy)
Mexico
Germany
Cayman Is
Cuba
Little Swan | (Honduras)
Cayman Is
Haiti, Dominican Republic
Haiti, Dominican Republic
Martinique
Saint Lucia
Galapagos (Ecuador
Galapagos (Ecuador
Galapagos (Ecuador
Galapagos (Ecuador
Species Extinction
LAST POSSIBLE
RECORDED CAUSE
1956 B
1850-1860 B,C/D
1726 =
1907 A,B,C/D
1935 A,B,C
1931
1890
1927 Cc
1964 c,D
1875 c
1907 c,D
1935
1931 A.C
1934 E
1888
1964
19th C
1968
1960s
o000
18th C
1950s
1902
19th C
1. Biological Diversity
Table 16.4 Animal species extinct since circa 1600 (continued)
SPECIES ENGLISH NAME DISTRIBUTION LAST POSSIBLE
RECORDED CAUSE
MAMMALS (continued)
Oryzomys victus St Vincent Rice Rat Saint Vincent 1897
* Peromyscus pembertoni Pemberton’s Deer Mouse Mexico
Family Echimyidae
# Boromys offella Cuba
# Boromys torrei Cuba
# Brotomys voratus Haiti, Dominican Republic
Family Muridae
Conilurus albipes Rabbit—eared Tree—rat Australia 1875
* Crateromys paulus llin Bushy —tailed Cloud—rat Philippines
Leporillus apicalis Lesser Stick—nest Rat Australia 1933
* Notomys amplus Short—tailed Hopping—mouse Australia 1894
* Notomys longicaudatus Long-tailed Hopping—mouse Australia 1901
+ Notomys macrotis Big—eared Hopping—mouse Australia pre—1850
+ Notomys mordax Darling Downs Hopping—mouse Australia pre—1846
+ Pseudomys fieldi Alice Springs Mouse Australia 1895
+ Pseudomys gouldi Gould's Mouse Australia 1930
Rattus macleari Maclear's Rat Christmas | peter 1908
Rattus nativitatis Bulldog Rat Christmas | (Australia 1908
Order CARNIVORA
Family Canidae
Dusicyon australis Falkland Island Wolf Falklands Is 1876 E
Family Procyonidae
+ Procyon gloveralleni Barbados Racoon Barbados
Order PINNIPEDIA
Family Phocidae
Monachus tropicalis Caribbean Monk Seal Caribbean 1962 A
Order SIRENIA
Family Dugongidae
Hydrodamalis gigas Steller’s Sea Cow Bering Straits (‘USSR’) 1768 A
Order PERISSODACTYLA
Family Equidae
Equus quagga Quagga South Africa 1883 AE
Order ARTIODACTYLA
Family Bovidae
Gazella rufina Red Gazelle Algeria? 19th C A
Hippotragus leucophaeus Bluebuck South Africa 1800 E
Family Cervidae
Cervus schomburgki Schomburgk's Deer Thailand 1932 A
d indicates species generally regarded as extinct but for which there may still be some chance of survival. + indicates taxa which may be
conspecific with extant forms. # indicates species known from post—Columbian (i.e. post 1500) deposits in the Caribbean; some may have become extinct
before 1600. . indicates species last recorded from Rapa in 1934, and which were considered likely to become rapidly extinct. x indicates species recorded
from subfossil deposits in the Mascarenes which are considered very likely to have become extinct following settlement in 1723 although may possibly have
become extinct earlier.
Possible causes column’: A Hunting (includes for food, skin, sport, live trade, feathers); B Direct habitat alteration by man; C Introduced predators
(e.g. cats, rats, mustelids, mongooses, snails, monkeys); (C/D predators or others not specified); D Other introduced animals (e.g. goats, rabbit, pigs);
E Destroyed as a pest species; F Introduced disease; G Indirect effects; H Natural Causes; — causes uncertain.
Note: The proceedings of a symposium entitled St Helena Natural Treasury (Edited by P. Pearce—Kelly and Q.C.B. Cronk, published by the Zoological
Society of London, 1990) were procured too late to include data in these lists. An additional eight extinct endemic bird species are listed from that island, six
of which should be included in our analysis. They are thought to have become extinct as a result of the human discovery of the island in 1502, and should
therefore be included in the same sort of category as those species recovered from post—Columbian deposits in the Caribbean (i.e. those marked #). The
report would also seem to indicate that it may be premature to declare the two insects Labidura herculeana and Aplothorax burchelli extinct, and they
should perhaps be excluded from this list at present. The effect these additions and changes have on the graphs and maps should be borne in mind,
especially the increase in early island bird extinctions.
Source: compiled from multiple sources; details available from WCMC. Most bird data compiled by A Stattersfield, and kindly made available by the
International Council for Bird Preservation. Mollusc data assembled by Sue Wells with the assistance of members of the SSC Mollusc Specialist Group and
other malacologists.
214
Table 16.5 Extinct higher plant taxa*
MAJOR GROUP (DIVISION)
FAMILY
TAXON
Fern Allies
Lycopodiaceae
Huperzia nutans Brackenr.
Selaginellaceae
Selaginella orizabensis Hieron.
Isoetaceae
/soetes dixitii Shende
/soetes sampathkumarnii L.N. Rao
Aspidiaceae
Diplazium laffanianum (Baker) C.Chr.
Dryopteris speluncae (L.) Underwood
Lastreopsis wattii (Beddome) Tagawa
Aspleniaceae
Asplenium fragile K. Pres| var. insularis C. Morton
Asplenium leucostegioides Baker
Diellia manii
Diellia unisora Wagner
Blechnaceae
Doodia /yoni Degener
Marsileaceae
Marsilea paradoxa Diels
Ophioglossaceae
Botrychium subbifoliatum Brackenr.
Thelypteridaceae
Christella altissima Holttum
Thelypteris macilenta E. St. John
Gymnosperms
Zamiaceae
Encephalartos woodii Sander
Zamia monticola Chamberlain
Dicots
Acanthaceae
Dicliptera abuensis Blatter
Dicliptera falcata (Lam.) Bosser & Heine
Hypoestes inconspicua Balf. f.
Hypoestes rodriguesiana Balf. f.
Hypoestes serpens R. Br.
Justicia brachystachya Thouars ex Schultz
Justicia eranthemoides F. Muell.
Justica psychotrioides Thouars ex Schultz
Aizoaceae
Gibbaeum esterhuyseniae L. Bolus
Trianthema cypseloides (Fenzl) Benth.
Amaranthaceae
Achyranthes atollensis St. John
Achyranthes mutica A. Gray ex H. Mann
Amaranthus mentegazzianus Passer.
Blutaparon rigidum (Robinson & Greenman) Mears
Ptilotus caespitulosus F. Muell.
Ptilotus extenuatus Benl
Ptilotus fasciculatus Fitzg.
Ptilotus pyramidatus (Moq.) F. Muell.
Anacardiaceae
Buchanania mangoides F. Muell.
Aquifoliaceae
Nex ternatiflora (C. Wright) R.A. Howard
Asclepiadaceae
Caralluma arenicola N.E. Brown
Marsdenia coronata Benth.
Marsdenia tubulosa F. Muell.
Matelea balbisii (Dcne.) Woods.
Matelea radiata Correll
Begoniaceae
Begonia cowellii Nash
Begonia opuliflora Putz.
Boraginaceae
Cryptantha aperta (Eastw.) Payson
Cryptantha insolita (J.F. Macbr.) Payson
Heliotropium muticum Domin
COMMON NAME
makou
Edward’s maiden fern
Balbis’ milkvine
Falfurrias Anglepod
Grand Junction cat's-eye
unusual cat’s-eye
Species Extinction
HISTORIC RANGE
United States - Hawaii
Mexico - Veracruz
India - Maharashtra State
India - Karnataka State
Bermuda
Bermuda
India - Manipur State
United States - Hawaii
United States - Hawaii
United States - Hawaii
United States - Hawaii
United States - Hawaii
Australia - Western Australia
United States - Hawaii
South Africa - Natal
United States - Florida
South Africa - Natal
Mexico
India - Rajasthan State
Mauritius
Mauritius - Rodrigues
Mauritius - Rodrigues
Mauritius
Mauritius
Australia - New South Wales
Mauritius
South Africa - Cape Province
Australia - New South Wales
United States - Hawaii
United States - Hawaii
Argentina
Ecuador - Galapagos
Australia - Western Australia
Australia - New South Wales
Australia - Western Australia
Australia - Western Australia
Australia - Queensland
Cuba
South Africa - Cape Province
Australia - Queensland
Australia - NSW - Lord Howe Island
United States - Arizona
United States - Texas
Cuba
Panama
United States - Colorado
United States - Nevada
Australia - Western Australia
1. Biological Diversity
Table 16.5 _—_ Extinct higher plant taxa*
MAJOR GROUP (DIVISION)
FAMILY
TAXON COMMON NAME HISTORIC RANGE
Heliotropium pannifolium Burchell ex Hemsley St Helena
Lindelofia angustifolia (Schrenk) A. Brand. former Union of Soviet Socialist
Republics
Myosotis petiolata Hook.f. var. pottsiana L. Moore New Zealand - North Island
Onosma affine Hausskn. ex H. Rield Turkey
Onosma discedens Hausskn. ex Bornm. Turkey
United States - California
United States - Oregon
Australia - South Australia
Plagiobothrys diffusus (Greene) |.M. Johnston San Francisco popcornflower
Plagiobothrys lamprocarpus (Piper) |.M. Johnston popcomflower
Plagiobothrys orthostatus J. Black
Bruniaceae
Staavia trichotoma (Thunb.) Pillans
Thamnea depressa Oliver
Thamnea uniflora Solander ex Brongn.
South Africa - Cape Province
South Africa - Cape Province
South Africa - Cape Province
Cactaceae
Hylocereus cubensis Britton & Rose Cuba
Leptocereus wrightii Leon Cuba
Lobivia vatteri Krainz Argentina
Opuntia lindheimeri Engelmann var. linguiformis
(Griffiths) L. Benson
United States - Texas
Pyrrhocactus aricensis Ritt. Chile
Pyrrhocactus longirama Ritt. Chile
Pyrrhocactus nuda Ritt. Chile
Pyrrhocactus occultus Ritt. Chile
Campanulaceae
Campanula oligosperma Damboldt Turkey
Clermontia multiflora Hillebrand
Cyanea arborea (H. Mann) Hillebrand var. arborea
Cyanea asplenifolia (H. Mann) Hillebrand
Cyanea comata Hillebrand
Cyanea dunbarii Rock
Cyanea giffardii Rock
Cyanea glabra (F. Wimmer) St. John
Cyanea grimesiana Gaudich. ssp. cylindrocalyx
(Rock) Lammers
Cyanea linearifolia Rock
Cyanea longissima (Rock) St. John
Cyanea obtusa (A. Gray) Hillebrand
Cyanea pohaku Lammers
Cyanea procera Hillebrand
Cyanea profuga C. Forbes
Cyanea pycnocarpa (Hillebrand) F.E. Wimmer
Cyanea quercifolia (Hillebrand) F.E. Wimmer var.
quercifolia
Cyanea recta (Wawra) Hillebrand
Cyanea scabra Hillebrand var. /ongissima Rock
Cyanea undulata C. Forbes
Delissea fallax Hillebrand
Delissea /aciniata Hillebrand var. /aciniata
Delissea /auliiana Lammers
Delissea parviflora Hillebrand
Delissea rivularis (Rock) F.E. Wimmer
Delissea sinuata Hillebrand ssp. /anaiensis (Rock)
Lammers
Delissea sinuata Hillebrand var. sinuata
Delissea undulata Gaudich.
Lobelia monostachya (Rock) Lammers
Lobelia remyi Rock
Rollandia parvifolia C. Forbes
Rollandia purpurellifolia Rock
Wahlenbergia burchellii A.DC.
Wehlenbergia roxburghii A.DC.
Wahlenbergia saxifragoides VV. Brehm.
Caryophyllaceae
Alsinidendron viscosum (H. Mann) Sherff
Schiedea amplexicaulis H. Mann
Schiedea helleri Sherff
Schiedea implexa (Hillebrand) Sherff
Schiedea spergulina A. Gray var. /leiopoda Sherff
Schiedea stellarioides H. Mann var. stellarioides
Silene cryptopetala Hillebrand
Silene oligotricha Huber-Mor.
Silene rectiramea Robinson
spleenwort-leaved cyanea
smooth cyanea
cut-leaf delissea
small-flowered delissea
wavy-leaf delissea
undulata delissea
laulihilihi; kawelu; ma’‘oli‘oli
216
United States - Hawaii
United States - Hawaii
United States - Hawaii
United States - Hawaii
United States - Hawaii
United States - Hawaii
United States - Hawaii
United States - Hawaii
United States - Hawaii
United States - Hawaii
United States - Hawaii
United States - Hawaii
United States - Hawaii
United States - Hawaii
United States - Hawaii
United States - Hawaii
United States - Hawaii
United States - Hawaii
United States - Hawaii
United States - Hawaii
United States - Hawaii
United States - Hawaii
United States - Hawaii
United States - Hawaii
United States - Hawaii
United States - Hawaii
United States - Hawaii
United States - Hawaii
United States - Hawaii
United States - Hawaii
United States - Hawaii
St Helena
St Helena
South Africa - Cape Province
United States - Hawaii
United States - Hawaii
United States - Hawaii
United States - Hawaii
United States - Hawaii
United States - Hawaii
United States - Hawaii
Turkey
United States - Arizona
Table 16.5
MAJOR GROUP (DIVISION)
FAMILY
TAXON
Stellaria elatinoides Hook. f.
Celastraceae
Hexaspora pubescens C. White
Maytenus lineata C. Wright
Chenopodiaceae
Hemichroa mesembryanthema F. Muell.
Sclerolaena ramsayae (Willis) A.J. Scott
Suaeda duripes |.M. Johnston
Compositae
Abrotanella rhynchocarpa Balf. f.
Acanthocladium dockeri F. Muell.
Argyroxiphium virescens Hillebrand var. virescens
Artemisia insipida Vill.
Brachycome muelleri Sonder
Calocephalus globosus M. Scott & Hutch.
Cirsium toyoshimae Koidz.
Commidendrum rotundifolium (Roxb.) DC.
Crepidiastrum ameristophyllum (Koidz.) Nakai
Crepidiastrum grandicollum (Koidz.) Nakai
Erigeron perglaber Blake
Felicia annectens (Harvey) Grau
Helianthus praetermissus E. Watson
Helichrysum oligochaetum F. Muell.
Helichrysum selaginoides (Sonder & F. Muell.)
Benth.
Helichrysum spiceri F. Muell.
Helipterum guilfoylei Ewart
Hemizonia mohavensis Keck
Leptorhynchos gatesii (Williamson) J.H. Willis
Lipochaeta bryanii Sherff
Lipochaeta ovata R. Gardner
Lipochaeta perdita Sherff
Marasmodes undulata Compton
Olearia arida Pritzel
Olearia flocktoniae Maiden & E. Betcke
Olearia oliganthema Benth.
Osteospermum hirsutum Thunb.
Perityle inyoensis (Ferris) A. Powell
Perityle villosa (Blake) Shinn.
Senecio behrianus Sonder & F. Muell.
Senecio georgianus DC.
Senecio /aticostatus Belcher
Senecio sandwicensis Less.
Solidago porteri Small
Tetramolopium arenarium (A. Gray) Hillebrand var.
arenarium
Tetramolopium arenarium (A. Gray) Hillebrand var.
confertum Sherff
Tetramolopium arenarium (A. Gray) Hillebrand ssp.
laxum Lowrey
Tetramolopium capillare (Gaudich.) H. St. John
Tetramolopium consanguineum (A. Gray)
Hillebrand ssp. consanguineum
Tetramolopium conyzoides (A. Gray) Hillebrand
Tetramolopium lepidotum Less. ssp. arbusculum
(A. Gray) T.K. Lowrey
Tetramolopium tenerrimum (Less.) Nees
Tracyina rostrata Blake
Vernonia africana (Sonder) Druce
Crassulaceae
Crassula alcicornis Schonl.
Crassula subulata Hermann var. hispida (Schonl. &
E.G. Baker) Toelken
Echeveria faui Moran & Meyran
Sedum pinetorum Brandegee
Sedum polystriatum R.T. Clausen
Tacitus bellus Moran & Meyran
Cruciferae
Extinct higher plant taxa*
COMMON NAME
hardtoe seepweed
greensword
Mojave tarweed; Mojave tarplant
ko’oko’olau; nehe
Inyo laphamia
Hanaupah laphamia
Porter’s goldenrod
showy indian clover
Pine City stonecrop
Species Extinction
HISTORIC RANGE
New Zealand
Australia - Queensland
Cuba
Australia - South Australia
Australia - Victoria
United States - Texas
Mauritius - Rodrigues
Australia - New South Wales, South
Australia
United States - Hawaii
France
Australia - South Australia
Australia - Western Australia
Japan
St Helena
Japan - Ogasawara-Shoto
Japan - Ogasawara-Shoto
United States - Arizona
South Africa - Cape Province
United States - New Mexico
Australia - Western Australia
Australia - Tasmania
Australia - Tasmania
Australia - Western Australia
United States - California
Australia - Victoria
United States - Hawaii
United States - Hawaii
United States - Hawaii
South Africa - Cape Province
Australia - South Australia, Western
Australia
Australia - New South Wales
Australia - New South Wales
South Africa - Cape Province
United States - California
United States - California
Australia - New South Wales, South
Australia, Victoria
Australia - South Australia, Victoria,
Western Australia
Australia - Victoria
United States - Hawaii
United States - Georgia, North Carolina
United States - Hawaii
United States - Hawaii
United States - Hawaii
United States - Hawaii
United States - Hawaii
United States - Hawaii
United States - Hawaii
United States - Hawaii
United States - California
South Africa - Natal
South Africa - Cape Province
South Africa - Cape Province
Mexico - Oaxaca
United States - California
Turkey
Mexico - Chihuahua
1. Biological Diversity
Table 16.5
MAJOR GROUP (DIVISION)
FAMILY
TAXON
Ballantinia antipoda (F. Muell.) E. Shaw
Caulanthus lemmonii
Diplotaxis siettiana Maire
Hutchinsia tasmanica Hook.
/satis arnoldiana N. Busch.
Lepidium drummondii Thell.
Lepidium merrallii F. Muell.
Lepidium obtusatum Kirk
Lepidium peregrinum Thell.
Menkea draboides (Hook.f.) Benth.
Phlegmatospermum drummondii (Benth.) O.
Schultz
Phlegmatospermum richardsii (F. Muell.) E. Shaw
Rorippa coloradensis Stuckey
Stroganowia sagittata Karelin & Kir.
Tropidocarpum capparideum Greene
Cucurbitaceae
Benincasa hispida (Thunb.) Cogn.
Sicyos hillebrandii H. St. John
Sicyos villosa Hook. f.
Dicrastylidaceae
Dicrastylis morrisonii Munir
Dilleniaceae
Hibbertia sargentii S. Moore
Epacridaceae
Andersonia bifida L. Watson
Andersonia longifolia (Benth.) L. Watson
Choristemon humilis Williamson
Coleanthera coelophylla (DC.) Benth.
Coleanthera virgata Stschegl.
Leucopogon cryptanthus Benth.
Leucopogon pogonocalyx Benth.
Ericaceae
Arctostaphylos uva-ursi (L.) Sprengel var.
franciscana (Eastw.) Roof
Arctostaphylos uva-ursi (L.) Sprengel var.
leobreweri Roof
Erica acockii Compton
Frica bolusiae Salter
Erica jasminiflora Salisb.
Erica pyramidalis Solander
Frica turgida Salisb.
Frica verticillata Bergius
Rhododendron mucronulatum Turcz. var. albiflora
Nakai
Erythroxylaceae
Erythroxylum echinodendron Ekman
Euphorbiaceae
Acalypha rubra Roxb.
Amperea protensa Nees
Beyeria cygnorum (Muell. Arg.) Benth.
Beyeria lepidopetala F. Muell.
Bonania myrcifolia (Griseb.) Benth. & Hook.
Chamaesyce celastroides (Boiss.) Croizat &
Degener var. tomentella
Claoxylon grandifolium (Poiret) Muell. Arg.
Cnidoscolus fragrans (H.B.K.) Pohl
Croton magneticus Airy Shaw
Euphorbia carissoides Bailey
Euphorbia daphnoides Balf. f.
Pseudanthus nematophorus F. Muell.
Fagaceae
Quercus boytoni Beadle
Frankeniaceae
Frankenia conferta Diels
Frankenia decurrens Summerh.
Frankenia parvula Turcz.
Gesneriaceae
Cyrtandra cyaneoides Rock
Extinct higher plant taxa*
COMMON NAME
Colorado watercress
caper-fruited tropidocarpum
‘akoko; koko; ‘ekoko; kokomalei
Boyton’‘s sand post oak
HISTORIC RANGE
Australia - Tasmania, Victoria
United States - Arizona
Spain
Australia - Tasmania
former Union of Soviet Socialist
Republics
Australia - Western Australia
Australia - Western Australia
NEW ZEALAND - North Island
Australia - New South Wales
Australia - Western Australia
Australia - Western Australia
Australia - South Australia, Western
Australia
United States - Colorado
Asiatic former Union of Soviet Socialist
Republics
United States - California
Australia - Queensland
United States - Hawaii
Ecuador - Galapagos
Australia - Western Australia
Australia - Western Australia
Australia - Western Australia
Australia - Western Australia
Australia - Victoria
Australia - Western Australia
Australia - Western Australia
Australia - Western Australia
Australia - Western Australia
United States - California
United States - California
South Africa - Cape Province
South Africa - Cape Province
South Africa - Cape Province
South Africa - Cape Province
South Africa - Cape Province
South Africa - Cape Province
Republic of Korea
Cuba
St Helena
Australia - Western Australia
Australia - Western Australia
Australia - Western Australia
Cuba
United States - Hawaii
Mauritius; France - Reunion
Cuba
Australia - Queensland
Australia - Queensland
Mauritius - Rodrigues
Australia - Western Australia
United States - Texas
Australia - Western Australia
Australia - Western Australia
Australia - Western Australia
United States - Hawaii
Species Extinction
Table 16.5 Extinct higher plant taxa*
MAJOR GROUP (DIVISION)
FAMILY
TAXON
Cyrtandra gracilis Hillebrand
Cyrtandra honolulensis Wawra
Cyrtandra kohalae Rock
Cyrtandra olona C. Forbes
Cyrtandra pickeringii A. Gray var. pickeringii
Cyrtandra waiolani Wawra var. capitata Hillebrand
Cyrtandra waiolani Wawra var. waiolani
Goodeniaceae
Dampiera helmsii Krause
Dampiera humilis (F. Muell.) E. Pritzel
Dampiera rupicola S. Moore
Goodenia clementii Krause
Scaevola attenuata R. Br.
Scaevola macrophylla (Vriese) Benth.
Scaevola oldfieldii F. Muell.
Verreauxia verreauxii (Vriese) Carolin
Grossulariaceae
Ribes kolymense (Trautv.) Komarov ex Pojark
Haloragaceae
Gonocarpus intricatus (Benth.) Orch.
Haloragis stricta R. Br.
Haloragis tenuifolia Benth.
Haloragodendron lucasii (Maiden & E. Betch) Orch.
Meziella trifida (Nees) Schindler
Hydrophyllaceae
Phacelia amabilis Constance
Phacelia cinerea Eastw.
Phacelia nevadensis J. Howell
Labiatae
Haplostachys bryanii Sherff var. bryanii
Haplostachys linearifolia (Drake) Sherff var.
linearifolia
Haplostachys munroi C. Forbes
Haplostachys truncata (A. Gray) Hillebrand
Hemigenia exilis S. Moore
Hemigenia obtusa Benth.
Hemigenia pimelifolia F. Muell.
Hemigenia podalyrina F. Muell.
Hemigenia ramosissima Benth.
Hemigenia tysoni F. Muell.
Hemigenia tysonii F. Muell.
Microcorys pimeloides F. Muell.
Monardella leucocephala A. Gray
Monardella pringlei A. Gray
Phyllostegia brevidens A. Gray var. brevidens
Phyllostegia hillebrandii Mann ex Hillebrand
Phyllostegia immunata (Sherff) St. John
Phyllostegia knudsenii Hillebrand
Phyllostegia rockii Sherff
Phyllostegia variabilis Bitter
Phyllostegia wawrana Sherff
Prostanthera staurophylia F. Muell.
Pycnanthemum monotrichum Fern.
Stenogyne cinerea Hillebrand
Stenogyne haliakalae Wawra
Stenogyne oxygona Degener & Sherff
Stenogyne viridis Hillebrand
Teucrium leucophyllum Montbret & Aucher ex
Bentham
Thymus oehmianus Ronn. & Soska
Lauraceae
Cassytha pedicellosa J.Z. Webb
Leguminosae
Acacia forrestiana E. Pritzel
Acacia murruboensis Maiden & Blakely
Acacia prismifolia E. Pritzel
Acacia vassalii Maslin
Aspalathus variegata Ecklon & Zeyher
Astragalus pseudocylindraceus Bornm.
Astragalus robbinsii (Oakes) A. Gray var. robbinsii
COMMON NAME
ha’‘iwale; kanawao ke’oke’o
Saline Valley phacelia
ashy phacelia
Nevada phacelia
Merced monardella
Pringle monardella
mountain mint
HISTORIC RANGE
United States - Hawaii
United States - Hawaii
United States - Hawaii
United States - Hawaii
United States - Hawaii
United States - Hawaii
United States - Hawaii
Australia - Western Australia
Australia - Western Australia
Australia - Western Australia
Australia - Western Australia
Australia - Western Australia
Australia - Western Australia
Australia - Western Australia
Australia - Western Australia
former Union of Soviet Socialist
Republics
Australia - Western Australia
Australia - New South Wales,
Queensland
Australia - Western Australia
Australia - New South Wales
Australia - Western Australia
United States - California
United States - California
United States - Nevada
United States - Hawaii
United States - Hawaii
United States - Hawaii
United States - Hawaii
Australia - Western Australia
Australia - Western Australia
Australia - Western Australia
Australia - Western Australia
Australia - Western Australia
Australia - Western Australia
Australia - Western Australia
Australia - Western Australia
United States - California
United States - California
United States - Hawaii
United States - Hawaii
United States - Hawaii
United States - Hawaii
United States - Hawaii
United States - Hawaii
United States - Hawaii
Australia - New South Wales
United States - Virginia
United States - Hawaii
United States - Hawaii
United States - Hawaii
United States - Hawaii
Turkey
Yugoslavia
Australia - Tasmania
Australia - Western Australia
Australia - New South Wales
Australia - Western Australia
Australia - Western Australia
South Africa - Cape Province
Turkey
United States - Vermont
1. Biological Diversity
Table 16.5 —_ Extinct higher plant taxa*
MAJOR GROUP (DIVISION)
FAMILY
TAXON
Chorizema varium Benth.
Crotalaria urbaniana Senn
Gastrolobium crispifolium Domin
Genista melia Boiss.
Jacksonia hemisericea D. Herbert
Lathyrus dominianus Litv.
Lupinus sublanatus Eastw.
Mirbelia densiflora C. Gardner
Onobrychis aliacmonia Reich. f.
Orbexilum macrophyllum Rydb.
Oxylobium acutum (Benth.) Benth.
Phyllota gracilis Turcz.
Psoralea macrophylla Rowlee ex Small
Psoralea stipulata Torrey ex A. Gray
Pultenaea pauciflora M. Scott
Sophora toromiro (Philippi) Skottsb.
Streblorrhiza speciosa Endl.
Taverniera sericophylia Balf. f.
Tephrosia kassasi Boulos
Tetragonolobus wiedemannii Boiss.
Trifolium amoenum E. Greene
Vicia dennesiana H.C. Watson
Lentibulariaceae
Utricularia mairii Cheeseman
Loasaceae
Mentzelia nitens Greene var. /eptocaulis J. Darl.
Loganiaceae
Mitrasacme palustris W. Fitzg.
Loranthaceae
Dendrophthora terminalis Kuljt
Psittacanthus nudus (A. Molina) Kuijt & Feuer
Trilepidea adamsii (Cheeseman) Tieghem
Malvaceae
Abutilon mauritianum (Jacq.) Medik.
Anisodontea alexandri (Baker f.) Bates
Hibiscadelphus bombycinus C. Forbes
Hibiscadelphus crucibracteatus Hobdy
Hibiscadelphus wilderianus Rock
Hibiscus nelsonii Rose & Standley
Kokia lanceolata Lewton
Malacothamnus abbottii (Eastw.) Kearney
Malacothamnus mendocinensis (Eastw.) Kearney
Sida pritzellii C. Gardner
Sidalcea keckii Wiggins
Sphaeralcea procera C.L. Porter
Menispermaceae
Hyperbaena obovata Urban
Menyanthaceae
Nymphoides stygia (J. Black) H. Eichler
Myoporaceae
Eremophila adenotricha F. Muell.
Eremophila scaberu/a Fitzg.
Myrsinaceae
Badula ovalifolia A.DC.
Myrsine mezii Hosaka
Myrtaceae
Calothamnus blepharantherus F. Muell.
Hypocalymma longifolium F. Muell.
Melaleuca arenaria C. Gardner
Melaleuca arenicola S. Moore
Melaleuca graminea S. Moore
Monimiastrum fasciculatum Gueho & A.J. Scott
Syzygium balfourii (Baker) Gueho & A.J. Scott
Verticordia carinata Turcz.
Nyctaginaceae
Pisonia floridena Britton
Ochnaceae
Ouratea alternifolia (A. Rich.) M. Gomez
Oleaceae
Hesperelaea palmeri A. Gray
Onagraceae
COMMON NAME
Santa Catalina Island desert-thorn
bigleaf scurpea
scurf-pea
toromiro
showy indian clover
Abbott’s bush-mallow
Mendocino bush-mallow
Keck sidalcea; Keck checker-mallow
Luna County globemallow
rock dey devil’s-claws
220
HISTORIC RANGE
Australia - Western Australia
Cuba
Australia - Western Australia
Greece
Australia - Western Australia
former Union of Soviet Socialist
Republics
United States - California
Australia - Western Australia
Greece
United States - North Carolina
Australia - Western Australia
Australia - Western Australia
United States - North Carolina
United States - Indiana, Kentucky
Australia - Western Australia
Chile - Easter Island
Australia - Norfolk Island
Democratic Yemen - Socotra
Egypt
Greece
United States - California
Portugal - Azores
New Zealand - North Island
United States - Arizona
Australia - Western Australia
Costa Rica
Honduras
New Zealand - North Island
Mauritius
South Africa - Cape Province
United States - Hawaii
United States - Hawaii
United States - Hawaii
Mexico
United States - Hawaii
United States - California
United States - Arkansas, California
Australia - Western Australia
United States - California
United States - New Mexico
Cuba
Australia - South Australia
Australia - Western Australia
Australia - Western Australia
France - Reunion
United States - Hawaii
Australia - Western Australia
Australia - Western Australia
Australia - Western Australia
Australia - Western Australia
Australia - Western Australia
Mauritius
Mauritius - Rodrigues
Australia - Western Australia
United States - Florida
Cuba
Mexico
Table 16.5
MAJOR GROUP (DIVISION)
FAMILY
TAXON
Clarkia mosquinii E. Small ssp. xerophila E. Small
Lopezia conjugens Brandegee
Lopezia sinaloensis Munz
Oenothera kleinii W.L. Wagner & S.W. Mill
Papaveraceae
Eschscholzia rhombipetala E. Greene
Penaeaceae
Stylapterus micranthus R. Dahlgren
Piperaceae
Peperomia degeneri Yuncker
Peperomia hirta Balf. f.
Peperomia rodriguezi Balf. f.
Peperomia rossii Rendle
Piumbaginaceae
Armeria arcuata Welw. ex Boiss. & Reuter
Polygalaceae
Comesperma lanceolatum Benth.
Comesperma rhadinocarpum F. Muell.
Polygonaceae
Eriogonum truncatum Torrey & A. Gray
Portulacaceae
Calandrinia composita Nees
Calandrinia dielsii Poelin.
Calandrinia feltonii Skottsb.
Calandrinia sphaerophylia J. Black
Primulaceae
Lysimachia forbesii Rock
Lysimachia minoricensis J.D. Rodriguez
Proteaceae
Grevillea batrachioides McGillivray
Grevillea divaricata R. Br.
Grevillea flexuosa (Lindley) Meissner
Grevillea scabra Meissner
Hakea crassinervia Meissner
Hakea pulvinifera L. Johnson
Hakea tamminensis C. Gardner
/sopogon uncinatus R. Br.
Leucadendron comosum (Thunb.) R. Br. ssp.
homoeophyllum (Meisn.) |. Williams
Leucadendron spirale (Salisb. ex Knight) |. Williams
Mimetes stokoei Phillips & Hutch.
Persoonia leucopogon S. Moore
Sorocephalus tenuifolius R. Br.
Triunia robusta (C. White) D. Foreman
Pyrolaceae
Pyrola oxypetala Austin
Rhamnaceae
Cryptandra tubulosa Fenzl.
Cryptandra uncinata Grun.
Spyridium kalganense Diels
Spyridium microcephalum (Turcz.) Benth.
Trymalium albicans (Steudel) Reisseck
Trymalium urceolare (F. Muell.) Diels
Rosaceae
Potentilla multijuga Lehm.
Rubiaceae
Danais corymbosa Balf. f.
Gaertnera calycina Bojer
Gaertnera crassiflora Bojer
Gaertnera longifolia Bojer var. pubescens Verdc.
Gaertnera quadriseta A.DC.
Hedyotis foliosa (Hillebrand) Fosb.
Oldenlandia adscensionis (DC.) Cronk
Oldenlandia polyclada (F.Muell.) F, Muell.
Oldentlandia sieberi Baker var. sieberi
Opercularia hirsuta F. Muell ex Benth.
Opercularia ocolytantha Diels.
Ophiorrhiza brunonis Wight & Arn.
Ophiorrhiza caudata C. Fischer
Extinct higher plant taxa*
COMMON NAME
Klein’s evening-primrose; Wolf Creek
evening-primrose
diamond-petaled; California poppy
Contra Costa eriogonum; Mt Diablo
buckwheat
sharp-petal wintergreen
Ballona cinquefoil
Species Extinction
HISTORIC RANGE
United States - California
Mexico
Mexico
United States - Colorado
United States - California
South Africa - Cape Province
United States - Hawaii
Mauritius - Rodrigues
Mauritius - Rodrigues
Australia - Christmas Island
Portugal
Australia - Western Australia
Australia - Western Australia
United States - California
Australia - Western Australia
Australia - Western Australia
Falkland Islands
Australia - South Australia
United States - Hawaii
Spain - Balearic Islands
Australia - Western Australia
Australia - New South Wales
Australia - Western Australia
Australia - Western Australia
Australia - Western Australia
Australia - New South Wales
Australia - Western Australia
Australia - Western Australia
South Africa - Cape Province
South Africa - Cape Province
South Africa - Cape Province
Australia - Western Australia
South Africa - Cape Province
Australia - Queensland
United States - New York
Australia - Western Australia
Australia - South Australia
Australia - Western Australia
Australia - Western Australia
Australia - Western Australia
Australia - Western Australia
United States - California
Mauritius - Rodrigues
Mauritius
Mauritius
Mauritius
Mauritius
United States - Hawaii
Ascension Island
Australia - Queensland
Mauritius
Australia - Western Australia
Australia - Western Australia
India - Karnataka State, Kerala State,
Tamil Nadu State
India - Kerala State
1. Biological Diversity
Table 16.5
MAJOR GROUP (DIVISION)
FAMILY
TAXON
Ophiorrhiza radicans Gardn.
Phyllacanthus grisebachianus Hook. f.
Psychotria banaona Urban
Pyrostria ferruginea Verdc.
Rondeletia odorata Jacq. var. breviflora Hook.
Wendlandia angustifolia Wight
Rutaceae
Acmadenia candida |. Williams
Agathosma orbicularis Bartl. & Wendl. f.
Eriostemon falcatus P.G. Wilson
Galipea ossana DC.
Kodalyodendron cubensis Borh. & Acuna
Melicope adscendens (St. John & Hume) T.
Hartley & B. Stone
Melicope ballouii (Rock) T. Hartley & B. Stone
Melicope degeneri (B. Stone) T. Hartley & B. Stone
Melicope lydgatei (Hillebrand) T. Hartley & B.
Stone
Melicope ovalis (St. John) T. Hartley & B. Stone
Melicope quadrangularis (St. John & E. Hume) T.
Hartley & B. Stone
Melicope reflexa (St. John) T. Hartley & B. Stone
Melicope wailauensis (St. John) T. Hartley & B.
Stone
Pelea fatuhivensis F. Brown
Pelea obovata H. St. John
Phebalium daviesii Hook. f.
Phebalium lachnaeoides Cunn.
Zanthoxylum leonis Alain
Zieria adenophora Blakely
Santalaceae
Leptomeria dielsiana Pilger
Santalum fernandezianum F. Philippi
Sapindaceae
Euchorium cubense Ekman & Radlk.
Saxifragaceae
Astilbe crenatiloba (Britton) Small
Mitella prostrata Michaux
Saxifraga lactea Turcz.
Saxifraga oppositifolia L. ssp. amphibia (Sunderm.)
Braun-Blanquet
Scrophulariaceae
Agalinis stenophylia Pennell
Agalinis strictifolia Pennell
Castilleja cruenta Standley
Castilleja leschkeana J. Howell
Euphrasia arguta R. Br.
Euphrasia collina R.Br. ssp. muelleri (Wettst.)
Barker
Limosella pubiflora Pennell
Micranthemum micranthemoides (Nutt.) Wettst.
Mimulus brandegei Pennell
Mimulus clementii Domin
Mimulus traskiae A.L. Grant
Mimulus whipplei A.L. Grant
Orthocarpus pachystachyus A. Gray
Penstemon leptanthus Pennell
Penstemon pulchellus Lindl.
Seymeria havardii (Pennell) Stand
Verbascum calycosum Hausskn. & Murb.
Veronica euxina Turrill
Solanaceae
Lycium hassei Greene
Mellissia begonifolia (Roxb.) Hook. f.
Solanum bahamense L. var. rugelii D'Arcy
Solanum bauerianum Endl.
Extinct higher plant taxa*
COMMON NAME
crenate-lobed false goat’s-beard
narrow-leaved false foxglove
indian paintbrush
Point Reyes indian paintbrush
mudwort
Nuttall’s micranthemum
Santa Cruz Island monkey-flower
Santa Catalina monkey-flower
Whipple’s monkey-flower
shasta owl-clover
Sevier Plateau beardtongue
beautiful beardtongue
Eagle Pass seymeria
222
HISTORIC RANGE
India - Kerala State; Sri Lanka
Cuba
Cuba
Mauritius
Panama
India - Tamil Nadu State
South Africa - Cape Province
South Africa - Cape Province
Australia - Western Australia
Cuba
Cuba
United States - Hawaii
United States - Hawaii
United States - Hawaii
United States - Hawaii
United States - Hawaii
United States - Hawaii
United States - Hawaii
United States - Hawaii
France - French Polynesia - Marquesas
Is
United States - Hawaii
Australia - Tasmania
Australia - New South Wales
Cuba
Australia - New South Wales
Australia - Western Australia
Chile - Juan Fernandez
Cuba
United States - North Carolina,
Tennessee
Canada
former Union of Soviet Socialist
Republics
Germany; Switzerland
United States - Florida
United States - Louisiana
United States - Arizona
United States - California
Australia - New South Wales
Australia - New South Wales, South
Australia, Victoria
United States - Arizona
United States - Delaware, District of
Columbia, Maryland, New Jersey, New
York, Pennsylvania, Virginia
United States - California
Australia - Westem Australia
United States - California
United States - California
United States - California
United States - Utah
United States - New Mexico
United States - Texas
Turkey
Bulgaria
United States - California
St Helena
United States - Florida
Australia - NSW
Lord Howe Island
Australia - Norfolk Island
Table 16.5 Extinct higher plant taxa*
MAJOR GROUP (DIVISION)
FAMILY
TAXON COMMON NAME
Solanum cajamarcense Ochoa
Solanum nava Webb & Berthel.
Sterculiaceae
Astiria rosea Lindley
Sterculia khasiana Deb.
Trochetia parviflora Bojer ex Baker
Stylidiaceae
Stylidium merrallii (F.Muell.) E. Pritzel
Stylidium neglectum Mildbr.
Stylidium pseudocaespitosum Mildbr.
Styracaceae
Styrax portoricensis Krug & Urban
Theaceae
Franklinia alatamaha Marshall Franklin tree
Tremandraceae
Tetratheca deltoidea J. Thompson
Tetratheca elliptica J. Thompson
Tetratheca fasciculata J. Thompson
Tetratheca gunnii Hook. f.
Umbelliferae
Geocaryum bornmuelleri (Wolff) Engstr.
Geocaryum divaricatum (Boiss. & Orph.) Engstr.
Platysace dissecta (Benth.) Norman
Platysace eatoniae (F. Muell.) Norman
Trachymene croniniana F. Muell.
Xanthosia singuliflora F. Muell.
Zizia latifolia Small bristol golden alexanders
Urticaceae
Pilea thouarsiana Wedd.
Pilea trilobata (Poiret) Wedd.
Valerianaceae
Valeriana pratensis (Benth.) Steud.
Violaceae
/sodendrion pyrifolium A. Gray wahine noho kula
Viola cryana Gillot
Zygophyllaceae
Fagonia taeckholmiana Hadidi
Monocots
Amaryllidaceae
Caliphruria tenera Baker
Eucharis lehmannii Regel
Eucrosia mirabilis (Baker) Pax
Gethyllis esterhuyseniae
Gethyllis latifolia Masson ex Baker
Habranthus caeruleus (Griseb.) Traub
Mathieua galanthoides Klotzsch
Plagiolirion horsmannii Baker
Araceae
Anthurium leuconeurum Lemaire
Philodendron clementis C.Wright ex Griseb.
Burmanniaceae
Thismia americana N. Pfeiffer thismia
Centrolepidaceae
Centrolepis caespitosa D. Cooke
Commelinaceae
Sauvallea blainii C. Wright
Cyperaceae
Bulbostylis neglecta (Hemsley) C.B. Clarke
Carex aboriginum M.E. Jones Indian Valley sedge
Carex paupera Nelmes
Carex repanda C.B. Clarke
Cladium drummondii C.B. Clarke
Eleocharis bermudiana Britton
Fimbristylis compacta Turrill
Schoenus acuminatus R. Br.
Schoenus natans (F. Muell.) Benth.
Tetraria australiensis C.B. Clarke
Dioscoreaceae
Dioscorea pentaphylia L.
Rajania prestoniensis Knuth
Eriocaulaceae
Eriocaulon echinospermoideum Ruhl.
i)
N
w
Species Extinction
HISTORIC RANGE
Peru
Spain - Canary Islands
Mauritius
India - Meghalaya State
Mauritius
Australia - Western Australia
Australia - Western Australia
Australia - Western Australia
Puerto Rico
United States - Georgia
Australia - Western Australia
Australia - Western Australia
Australia - Western Australia
Australia - Tasmania
Greece
Greece
Australia - Western Australia
Australia - Western Australia
Australia - Western Australia
Australia - Western Australia
United States - Florida
Mauritius
Mauritius
Mexico
United States - Hawaii
France
Egypt
Colombia
Colombia
Ecuador
South Africa - Cape Province
South Africa - Cape Province
Argentina
Peru
Colombia
Mexico
Cuba
United States - Illinois
Australia - Western Australia
Cuba
St Helena
United States - Idaho
Australia - Victoria
India - Meghalaya State
Australia - Western Australia
Bermuda
Australia - Northern Territory
Australia - Western Australia
Australia - Western Australia
Australia - Western Australia
Australia - Queensland
Cuba
Cuba
1. Biological Diversity
Table 16.5 Extinct higher plant taxa*
MAJOR GROUP (DIVISION)
FAMILY
TAXON
Eriocaulon johnstonii Ruhl.
Eriocaulon minutessimum Ruhl.
Lachnocaulon cubense Ruhl.
Gramineae
Agrostis adamsonii Vick.
Agrostis limitanea J. Black
Bromus brachystachys Hommung
Bromus bromoideus (Lej.) Crepin
Bromus grossus Desf. ex DC.
Bromus interruptus (Hackel) Druce
Cenchrus agrimonioides Trin. var. laysanensis F.
Brown
Deyeuxia drummondii (Steudel) Vick.
Deyeuxia lawrencei Vick.
Digitaria pittieri (Hackel) Henrard
Dissanthelium californicum (Nutt.) Benth.
Eragrostis deflexa Hitchc.
Eragrostis fosbergii Whitney
Eragrostis hosakai Degener
Eragrostis mauiensis Hitchc.
Eragrostis rottleri Stapf
Eriochrysis rangacharii Fischer
Festuca benthamiana Vick.
Glyceria drumondii (Steudel) C.E. Hubb.
Heterachne baileyi C.E.Hubb.
Homopholis belsonii C.E.Hubb.
Hubbardia heptaneuron Bor
Paspalum amphicarpum Ekman
Paspalum jimenezii Chase
Plectrachne bromoides (F. Muell.) C.E. Hubb.
Poa manii Munroe ex Hillebrand
Poa mannii Munro
Streptochaeta angustifolia Soderstrom
Sucrea sampaiana (A. Hitch.) Soderstrom
Trisetum burnoufii Req. ex Parl.
Zea mays L. ssp. mexicana (Schrad.) Wilkes raza
durango
Hydatellaceae
Hydatella australis Diels
Hydatella leptogyne Diels
Hydrocharitaceae
Elodea linearis H. St. John
Elodea schweinitzii (Planchon) Casper
lridaceae
Gladiolus alatus L. var. algoensis Herb.
Hesperantha saldanhae P. Goldblatt
Iris antilibanotica Dinsm.
Iris damascena Mont.
Iris westii Dinsm.
Moraea incurva Lewis
Romulea papyracea W. Dod
Romulea sulphurea Beguinot
Sisyrinchium farwellii Bickn.
Sisyrinchium hastile Bickn.
Juncaceae
Juncus griscomii
Juncus oronensis Fern.
Juncus pervetus Fern.
Allium rouyi Gaut.
Calochortus indecorus Ownbey & M. Peck
Calochortus monanthus Ownbey
Dipcadi concanense (Dalz.) Baker
Dipcadi reidii Deb & Dasgupta
Ipheion tweedianum (Griseb.) Traub
Lachenalia mathewsii Barker
Smilax leptanthera Pennell
Tulipa sprengeri Baker
Urginea duthiae Adamson
Urginea ecklonii Baker
COMMON NAME
kamanomano; kumanomano
California dissanthelium
Pacific lovegrass
Fosberg’s lovegrass
Mann’‘s bluegrass
Nashville waterweed
Schweinitz’s waterweed
Farwell’s blue-eyed-grass
spear-like blue-eyed-grass
Griscom’s rush
Maine rush
Barnstable bog rush; old veteran rush
Sexton Mt mariposa-lily
Shasta River mariposa;
single-flowered mariposa lily
catbrier
224
HISTORIC RANGE
Mauritius
Cuba
Cuba
Australia - Victoria
Australia - South Australia
Germany
Belgium
Belgium; Luxembourg
United Kingdom
United States - Hawaii
Australia - Western Australia
Australia - Tasmania
Costa Rica
United States - California; Mexico
United States - Hawaii
United States - Hawaii
United States - Hawaii
United States - Hawaii
India - Tamil Nadu State
India - Tamil Nadu State
Australia - South Australia
Australia - Western Australia
Australia - Queensland
Australia - New South Wales,
Queensland
India - Karnataka State
Cuba
Costa Rica
Australia - Western Australia
United States - Hawaii
United States - Hawaii
Brazil
Brazil
France - Corsica
Mexico
Australia - Western Australia
Australia - Western Australia
United States - Tennessee
United States - Pennsylvania
South Africa - Cape Province
South Africa - Cape Province
Syria
Syria
Lebanon
South Africa - Cape Province
South Africa - Cape Province
South Africa - Cape Province
United States - Michigan
United States - Michigan
United States - Virginia
United States - Maine
United States - Massachusetts
Spain
United States - Oregon
United States - California
India
India
Argentina
South Africa - Cape Province
United States - Georgia
Turkey
South Africa - Cape Province
South Africa - Cape Province
Table 16.5 Extinct higher plant taxa*
MAJOR GROUP (DIVISION)
FAMILY
TAXON COMMON NAME
Urginea polyphylla Hook. f.
Orchidaceae
Palmae
Acrolophia ustulata Schlecther & Bolus
Caladenia atkinsonii Rodway
Caladenia pumila R. Rogers
Calanthe whiteana King & Pantl.
Corycium vestitum Sweet
Diuris fastidiosa R. Rogers
Paphiopedilum delenatii Guillaumin
Pleione lagenaria Lindley
Prasophyllum colemaniae R. Rogers
Prasophyllum subbisectum Nicholls
Satyrium guthriei Bolus
Triphora latifolia G. Luer nodding cape
Zeuxine boninensis Tuy
Acrocomia subinermis Leon ex L.H. Bailey
Corypha taliera Roxb.
Paschalococos disperta Dransfield
Pritchardiopsis jennencyi Becc.
Pandanaceae
Pandanus barklyi Belf. F. var. macrocarpus
Vaughan & Wiehe
Pandanus conglomeratus Balf. f.
Pandanus iceryi Horne ex Balf. f.
Pandanus incertus Vaughan & Wiehe
Pandanus macrostigma Martelli
Pandanus obsoletus Vaughan & Wiehe
Pandanus spathulatus Martelle
Restionaceae
Elegia extensa Pillans
Flegia fastigiata Mast.
Leptocarpus ramosissimus Pillans
Lepyrodia heleocharojdes Gilg
Restio chaunocoleus F. Muell
Tecophilaeaceae
Tecophilaea cyanocrocus Leybold
Zingiberaceae
Hedychium marginatum C.B. Clarke
Extinct Species
HISTORIC RANGE
India
South Africa - Cape Province
Australia - Tasmania
Australia - Victoria
India - Sikkim State
South Africa - Cape Province
Australia - Victoria
Vietnam
India - Meghalaya State
Australia - Victoria
Australia - Victoria
South Africa - Cape Province
United States - Florida
Japan - Ogasawara-Shoto
Cuba
India
Chile - Easter Island
France - New Caledonia
Mauritius
Mauritius
Mauritius
Mauritius
Mauritius
Mauritius
Mauritius
South Africa - Cape Province
South Africa - Cape Province
South Africa - Cape Province
Australia - Western Australia
Australia - Western Australia
Chile
India - Nagaland State
Notes: This list represents information available to WCMC in computerised form as of March 1992. It is intended to include species that are extinct
(or presumed extinct) in the wild, whether or not they are in cultivation. Several of these plants, such as Franklinia alatamaha, Paphiopedilum
delenatii and Tecophilaea cyanocrocus, are, in fact, well known in the horticultural trade. Others, such as Encephalartos woodii, are known only
from relatively few specimens, mostly held in botanic gardens. A few others have become extinct in the wild but have been reintroduced from
cultivated material grown in botanic gardens.
The information available is strongly biased geographically: many other species of higher plants have undoubtedly become extinct but lack of
country-based data prevents their inclusion here. Some of the species in this list are almost certainly still extant in remote, isolated areas; publication
of this list should stimulate searching for them.
* Includes some taxa below species level.
Notes for Table 16.6, overleaf: (1) Two amphibians (USA and Israel), one coral (Panama) and one mammal (Caribbean) are not included in this
table. (2) The above species may have lived in more than one country therefore total numbers do not necessarily agree with other tables. * indicates
islands which are not on the standard country list. They have been included separately because of the importance of islands when considering
extinctions.
225
1. Biological Diversity
Table 16.6 Known animal extinctions since c. 1600 by country
MOLLUSCS INSECTS FISHES REPTILES BIRDS MAMMALS TOTAL
ASIA
India
Indonesia
Nansei—shoto (Japan)*
Nepal
Ogasawara—shoto (Japan)*
-“~|O-$NN &
Philippines
Taiwan 1
Thailand 1
‘USSR’ it
Bering Straits (‘USSR’)* 1 1
EUROPE
Austia 1
Canary Islands (Spain)* 1
Corsica (France)* 1
Faeroe Islands 1
Germany 1 1
Iceland 1
Sardinia oe 1
United Kingdom
Yugoslavia 1
NORTH & CENTRAL AMERICA
Bahamas 1
Barbados 1 1
Canada 2 2
Cayman Is 1
Cuba 1
Ny + += F/0-NN
ae aioomana a
@ |
Dominican Rep
Greenland u
Guadalupe (Mexico)* 2
Guadeloupe 1 1
Guatemala 1
Haiti 6
Jamaica 2 1
Little Swan Island (Honduras)* 1
Martinique 3 1
Mexico 12 1 2
Navassa Island (USA)* 1
Saint Lucia 1
Saint Vincent and the Grenadines 1
United States 38 9 17 ‘ 4
Virgin Islands (US) 1
SOUTH AMERICA
Brazil 1
Colombia 1
Galapagos (Ecuador)* 1 4
Uruguay 1
OCEANIA
American Samoa 2
Australia 1 17
Bougainville (PNG)*
Chatham Island (NZ)* 1
Christmas Island (Australia)* 2
ea
aBasalRaawalsnnalmaan a
=u =
=
= o-
=
QinS-@n
Cook Islands 14
Fiji 1
French Polynesia 33
Guam 1
Hawaii (USA)* 29 42
BoB
Kangaroo Island (Australia)*
Lord Howe Island (Australia)* 1
Micronesia, Federated States of
New Caledonia 2
New Zealand 1 1 1
an
Es
oa
-“~|SCWON “(OA u
Norfolk Island (Australia)*
Palau 1
Phillip Island (Austraia)*
Solomon Islands
Stephens Island (NZ)* 1
arene
Tasmania (Australia)* 1
Wake Island (USA)* 1
Western Samoa 1
ANTARCTICA
Falkland Islands (Malvinas) & dependencies 1 1
AFRICA
Algeria 1 1
Ascension Island (UK)* 1 1
Cape Verde 1 1
Madagascar 3 3
Madeira (Portugal)* 14 14
44 [po--- a laAaoo =
Mauritius 23 1
Réunion 1
Rodrigues (Mauritius)* 6
Saint Helena 22 2 24
Seychelles 2 1 3
South Africa 2 2
Tristan da Cunha (UK)* 1 1
ena
_
=
=
_
a
226
Species Extinction
sejdeds jo sequiny
Figure 16.6 Known animal extinctions since c. 1600: Molluscs
0 —
S| 4009
IL
eiluopaje9 BUBI9H JUIES
, “Ee MON
f J Bs
| x a sen6upoy
oN __ sobedejey yi \ :
oe + x 6 |
xe eIseUuA|Od Bowes -
; ) = youaly uequewy
L ednojepend ie 3 cae ‘ ag ri
\ >
ee 2S
1. Biological Diversity
Figure 16.7 Known animal extinctions since c. 1600: Arthropods
se|oeds jo JeqUNN
(ZN) | weueyo
=
(eyeqsny)
| MOH Pso7
(ZN) | sueydeys | ray
2 puejeez MON
228
Species Extinction
Figure 16.8 Known animal extinctions since c. 1600: Fishes, reptiles and amphibians
seloeds jo JequINN,
sopeqieg mum <
ee Jenbiuuey Ss :
tig ena
A x
s| WBA SD) ome
ednojepeny =a”
| ESSeACN mmm
puejeez MaN
Ea
—
Uy
a
snguney
uojuney
sueiquduy =| |
sejgdey ES
seusiy [5]
1. Biological Diversity
Figure 16.9 Known animal extinctions since c. 1600: Birds
sejoeds jo JequiNnN
(ZN) | sueydeis
B puejesz MON
(eyeasny) snquney
(ZN) | wetpeyd | @MOH Pio}
3 aoe es (evensny)
. I ooueBuey ByUND ep uejysuy
i ray =a senBupoy 1. , 2)
v
AenBnuf) (eyensny) ie
$s] 4009 1 ditud
me OP IMIOHON
Pm os
ener won fT 5)
BOWES WOISOMA S| UOWOIOS = (DNd) ?
_ = allnuie6nog ty
Ws A)
-
9 wen seuiddiji
rere” = ms a : | 7, (ued)
“> (Coe) ~ -- : p —
SeWeYeG mam ednjepen5 (ueder) /~ 2 7 2a :
ojoys-esemeseBO 6
230
Species Extinction
Figure 16.10 Known animal extinctions since c. 1600: Mammals
seloeds jo equiny
e
sopeqieg mmm —.*
| Clon] JUIeG Sa 4 3
s| puepyle4
as)
enbiupey Se a
doy uediuilwi0g yi
H
(seunpuoH)
1 =| UBMs ef]
==
s| uewAe9
puejeez MeN
231
1. Biological Diversity
Table 16.7 Animal species surviving only in captivity
SPECIES
MOLLUSCS
Order STYLOMMATOPHORA
Family Partulidae
Partula spp.
Partula affinis
Partula aurantia
Partula clara
Partula hyalina
Partula mirabilis
Partula mooreana
Partula nodosa
Partula otaheitana
Partula suturalis
Partula taeniata
Partula tohiveana
FISHES
Order CYPRINODONTIFORMES
Family Cyprinodontidae **
Cyprinodon alvarezi
Megupsilon aporus
Family Poeciliidae
Xiphophorus couchianus
Family Goodeidae
Skiffia francesae
BIRDS
Order CICONIIFOMES
Family Ciconiidae
Gymnogyps californianus
Family Columbidae
Zenaida graysoni
MAMMALS
Order PERISSODACTYLA
Family Equidae
Equus ferus
Order ARTIODACTYLA
Family Bovidae
Bos taurus
ENGLISH NAME
Viviparous Tree Snails
Monterrey Platyfish
Golden Sawfin
Californian Condor
Socorro Dove
Wild Horse
Domestic cattle
NOTES
French Polynesia. Exterminated in wild after
introduction of Fuglandina rosea in 1977.
Various captive colonies around the world.
Population status information as at March
TEI Se
Tahiti. Functionally extinct, only 1 left alive.
Moorea. Functionally extinct, only 1 left alive.
Tahiti. Critical.
Tahiti. Increasing.
Moorea. Critical.
Moorea. Seriously declining.
Tahiti. Increasing.
Tahiti. Increasing but low numbers.
Moorea. Declining/stable.
Moorea. Increasing, good numbers.
Moorea. Increasing but all from 4 individuals.
Mexico. Last specimens removed from wild
February 1992.
Mexico. Last specimens removed from wild
February 1992; a number of captive
populations exist.
Mexico. Extinct in the wild in 1960s; three
captive populations.
Mexico; widespread in captivity.
USA. Last individual taken from wild 1987. 52
in captivity at end of 1991.
Socorro | (Mexico). Extinct post-1958. Large
captive populations.
China, Mongolia. Some disagreement on
taxonomic status. £. ferus gmelini, the Tarpan,
exterminated late 19th century. £ f.
przewalskii, Przewalski’s Horse survives in
zoos, last seen in wild in 1968.
Europe, North Africa and the Near East. The
Aurochs 8. t. primigenius, the wild ancestor
was exterminated in 1627.
Notes: * Reference: Partula 91, Proceedings of the Partula Propagation Group Meeting, 16 May 1991. Compiled by S. Tonge, WPT. ** Note
two further Cyprinodon species, Charco Azul and Charco Palma will probably also soon be extirpated in the wild (P. Loiselle, pers. comm.).
232
Species Extinction
Table 16.8 Animal species extirpated in wild and reintroduced
BIRDS
Order GRUIFORMES
Family Rallidae
Rallus owstoni
MAMMALS
Order CARNIVORA
Family Mustelidae
Mustela nigripes
Family Canidae
Canis rufus
Order ARTIODACTYLA
Family Bovidae
Bison bonasus
Oryx leucoryx
Family Cervidae
Elaphurus davidianus
Guam Rail
Black-footed Ferret
Red Wolf
Wisent
Arabian Oryx
Pére David’s Deer
Guam (USA). Extinct in wild 1985.
Reintroduced 1990/91.
USA. Last specimen taken from wild in 1987.
Reintroduced 1990/91.
USA. Extinct in wild 1980, reintroduced late
1980s.
Europe. Exterminated in wild by 1927.
Reintroduced to several locations.
Middle East. Last recorded in the wild in 1972.
Reintroduced in Oman in 1982.
Discovered in captivity in 1861. Now exists in
zoos worldwide. Reintroduced to China.
233
1. Biological Diversity
17. THREATENED SPECIES
A threatened species is one thought to be at significant risk
of extinction in the foreseeable future, because of stochastic
or deterministic factors affecting its populations, or by
virtue of inherent rarity. This convenient working definition
is deceptively simple; deciding what level of risk is
significant, and what part of the future is foreseeable, is
problematic.
WHAT IS A THREATENED SPECIES?
The growth in public awareness of the problem of depletion
and possible extinction of species is largely attributable to
the development of the Red Data Book (RDB) concept by
Sir Peter Scott during the 1960s. This involves an attempt
to categorise species at risk according to the severity of the
threats facing them and the estimated imminence of their
extinction. The RDBs were compiled on a global basis by
TUCN, so far as available information allowed, but the
concept was soon adopted at a national or sub-national level
in several countries. Attention also spread from the
terrestrial vertebrates, which were the principal focus of
early RDBs, to invertebrates and plants.
As the volume of information has increased, the traditional
Red Data Book approach, which included publication of a
range of data on each threatened species, has been to some
extent replaced by a direct listing of globally-threatened
species recognised by IUCN. The IUCN Red List of
Threatened Animals (IUCN, 1990, latest edition) is the only
accepted worldwide attempt to list threatened animal species
individually, and has provided the basis for the discussion
below.
The animals Red List has been compiled every two years
since 1986 by the World Conservation Monitoring Centre,
in collaboration with the JIUCN Species Survival
Commission network of Specialist Groups. The Red List is
based on information provided by numerous scientists,
naturalists and conservationists working in the field, much
of it collated by the IUCN SSC Specialist Groups. The
categorisation of threatened bird species is undertaken by
the International Council for Bird Preservation (ICBP).
Each species covered in the Red List is assigned a threat
category determined by a review of the factors affecting it
and the extent of the effect that these are having throughout
its range. Key factors examined include changes in
distribution or numbers, degree and type of threat, and
population biology. IUCN Red List categories are applied
to species on an international or global scale, and should
not be confused with the national threat categories assigned
to species by countries which have prepared Red Lists or
Red Data Books dealing with the status of species within
their own borders.
It is important to note that although the IUCN Red List is
a comprehensive global compendium of animal species
known to be threatened, many more species than those listed
will in fact be threatened. Those not listed fall into two
categories: first, and probably the largest number of
species, are those not yet described by science; and second,
the status of many described species has not been reviewed.
234
Birds have been comprehensively reviewed by ICBP; only
50% of mammal species, and probably less than 20% of
reptiles, 10% of amphibians and 5% of fish are estimated
to have been reviewed.
TUCN threat categories
The main IUCN threat categories currently used, together
with their definitions (as used in the Red Lists) are:
Extinct (Ex)
Species not definitely located in the wild during the past 50
years. On a few occasions, the category Ex? has been
assigned, denoting that it is virtually certain that the taxon
has recently become extinct.
Endangered (E)
Taxa in danger of extinction and whose survival is unlikely
if the causal factors continue to operate. Included are taxa
whose numbers have been reduced to a critical level or
whose habitats have been so drastically reduced that they
are deemed to be in immediate danger of extinction. Also
included are taxa that may now be extinct although they
have been seen in the wild in the past 50 years.
Vulnerable (V)
Taxa believed likely to move into the Endangered category
in the near future if the causal factors continue operating.
Included are taxa of which most or all the populations are
decreasing because of over-exploitation, extensive
destruction of habitat or other environmental disturbance;
taxa with populations that have been seriously depleted and
whose ultimate security has not been assured; and taxa with
populations that are still abundant but are under threat from
severe adverse factors throughout their range.
Rare (R)
Taxa with small world populations that are not at present
Endangered or Vulnerable but are at risk. These taxa are
usually localised within restricted geographical areas or
habitats or are thinly scattered over a more extensive range.
Indeterminate (1)
Taxa known to be Endangered, Vulnerable or Rare but
where there is not enough information to say which of the
three categories is appropriate.
Insufficiently Known (K)
Taxa that are suspected but not definitely known to belong
to any of the above categories, because of lack of
information.
The general term threatened is used to refer to a species
considered to belong to any one of the above categories.
The same definitions have been applied to plants, although
they have often been interpreted in a significantly different
manner, mainly because of biological differences between
animals and plants, and intermediate categories (e.g. Ex/E
or E/R) are also employed.
The definition and application of such status categories has
been a matter of some discussion, principally because they
provide such an important tool in assessing needs and
mobilising resources for conservation at the international,
national or sub-national level. In the opinion of many
scientists, the existing IUCN threat category definitions are
excessively subjective, and as a result categorisations made
by different authorities can vary and may not accurately
reflect real extinction risks. Mace and Lande (1991) have
recently proposed a new system based on quantitative (and
therefore theoretically objective) Population Viability
Analysis techniques.
The threats
Most of the causal factors currently threatening species are
anthropogenic in nature, i.e. induced or influenced by man.
These factors include:
e Habitat loss or modification, often associated with
habitat fragmentation. Causes include pastoral
development, cultivation and settlement, forestry
operations and plantations, fire, and pollution
Over-exploitation for commercial or subsistence reasons,
including meat, fur, hides, collection of live animals for
the pet trade and plants for the horticultural trade
Accidental or deliberate introduction of exotic species,
which may compete with, prey on or hybridise with
native species
Disturbance, persecution and uprooting, including
deliberate eradication of species considered to be pests
Incidental take, particularly the drowning of aquatic
reptiles and mammals in fishing nets
Disease, both exotic and endemic, exacerbated by the
presence of large numbers of domestic livestock or
introduced plant species
Figure 17.1 Analysis of threats: mammals
Habitat loss &
modification
60
so
40
30
20
Per cent of species affected
10
Other
Meat
Cultivation &
settlement
Pastoral development
Logging & plantations
Fur & hides
Exploitation
Live trade
Threats and classes of threat
235
Threatened Species
e Limited distribution, which may compound the effects of
other factors.
In the majority of cases individual species are faced by
several of these threats operating simultaneously, and it is
often difficult or impossible to identify with confidence the
primary cause of decline.
Some understanding of the relative importance of different
threat types, as measured by frequency of occurrence, can
be gained from an examination of threats facing the
mammals (excluding Cetacea) of Australasia and the
Americas (comprehensively reviewed by Thornback and
Jenkins, 1982), and those facing the birds of the world
(Diamond, 1987).
Of the 119 species of mammals from these continents
considered threatened, 75% (94) are threatened by more
than one factor, and of these, 27 face four or more threats.
The major category of threat, which affects 76% of species,
is habitat loss and modification (Fig. 17.1). This has a
variety of causes, of which the most frequent is cultivation
and settlement. Over-exploitation affects half the species,
the most significant cause being hunting for meat.
Introduced predators and competitors affect 18% of
threatened species. The most serious other factor is limited
distribution, which affects one quarter of species.
Fig. 17.2 compares the major threats affecting the birds of
the world with those affecting the mammals of Australasia
and the Americas. There is a high degree of similarity
between the two groups. Habitat destruction is the single
most important threat, affecting 60% of birds and 76% of
Others
Predators
Compet | tors
Limited distribution
Persecut Ion
Disturbance
Incidental take
Disease
1. Biological Diversity
Figure 17.2 Analysis of threats: mammals and birds
80
70
Per cent of threatened species
Habitat destruction
Hunting
Introduced species
International trade
Cof Australasia &
the Americas)
bees Mamma | Ss
Wetland drainage Incidental take
Pollution
Threat type
mammals. A major difference is that almost double the
number of mammals as birds are threatened by hunting
(54% versus 29%).
GLOBALLY THREATENED ANIMALS
Taxonomic distribution of threatened animals
The term ‘threatened’ in the following discussion refers to
taxa assigned a relevant status category by IUCN. In all,
some 4,452 animal species are listed as threatened in the
1990 Red List, or much less than 0.5% of the world’s
estimated total of well over 1.5 million described animal
species (Tables 17.1 and 17.2). Some species are also listed
in part only, i.e. one or more subspecies are included in the
Red List, but only full species are considered here.
The two classes with the greatest number of threatened
species are birds with 1,029 and insects with 1,083. Other
major listings include 507 mammals, 169 reptiles, 57
amphibians, 713 fish, 409 molluscs, 154 corals and
sponges, 139 annelid worms and 126 crustaceans. Clearly,
the number of threatened species in a taxonomic group is
not directly proportional to the overall number of species in
that group: some groups, particularly vertebrates, have
higher proportions listed as threatened than other groups.
The four major groups with the highest percentage of
threatened species are mammals (11.7% threatened), birds
(10.6%), fish (3.6%) and reptiles (3.5%). In comparison,
although a large number (1,083) of insects is listed, this
represents less than 0.15% of the world’s total. This
dichotomy between vertebrates and invertebrates becomes
even more extreme when Endangered species, the most
severely threatened category, are examined. Each of the
236
five vertebrate groupings have a higher percentage of listed
Endangered species than all of the invertebrate taxa added
together (Fig. 17.3).
Considering only the mammals among vertebrates, several
smaller orders have a very high proportion of threatened
species (Proboscidea with two out of two species, Sirenia
with four out of four species and Perissodactyla with 12 out
of 16 species). Among the larger orders, Primates,
Carnivora and Artiodactyla are the most threatened, with
respectively 53%, 32% and 31 % of their constituent species
listed. Although these three orders combined only contain
some 14.6% of the world’s mammal species, they account
for just under half of the listed threatened species and just
over half of the Endangered species.
To some extent, vertebrates may be more vulnerable to
extinction than invertebrates because they are typically
much larger and therefore require more resources and
larger ranges. On the other hand, many invertebrates have
an extremely small range, which would render them liable
to extinction by habitat loss. It seems reasonable to
conclude that the proportion of species in a group listed as
threatened reflects popular and scientific attention in
addition to biological reality.
Geopolitical distribution of threatened animals
Table 17.3 shows the geopolitical distribution of threatened
animal species according to the IUCN Red List (1990)
together with threatened plants; Table 17.4 shows a subset
of the animal data, with the countries listed in descending
order according to the number of threatened species in each
higher grouping. The top ten countries are listed for each
taxon.
Threatened Species
Table 17.1 IUCN Threatened Vertebrates (1990 Red List)
CLASS NUMBER OF SPECIES APPROXIMATE TOTAL
ORDER THREATENED ENDANGERED OF DESCRIBED SPECIES
MAMMALS 507 140 4,327
Monotremata 1 0 3
Marsupialia 25 6 282
Xenarthra 6 1 29
Insectivora 79 3 365
Scandentia {e) (0) 16
Dermoptera f0) 10) 2
Chiroptera 45 11 977
Primates 106 47 201
Pholidota fe) ie} 7
Lagomorpha 9 6 65
Macroscelidia 2 {e} 15
Rodentia 54 15 1,793
Cetacea 211 6 77
Carnivora 76 12 235
Pinnipedia 4 2 34
Sirenia (@) 5
Proboscidea 1 2
Perissodactyla 12 7 16
Hyracoidea 1 ie) 8
Tubulidentata {o} 0 1
Artiodactyla 60 23 194
BIRDS i 1,029 132 9,672
Struthioniformes 1 fe) 10
Tinamiformes 8 fe) 47
Sphenisciformes 3 (0) 17?
Podicipediformes 4 2 21?
Procellariformes 25 4 115?
Pelecaniformes 8 3 9?
Ciconiiformes 21 8 19?
Anseriformes 20 3 168
Falconiformes 45 6 311?
Galliformes 68 11 214
Gruiformes 51 a 196
Charadriiformes 31 4 350?
Columbiformes 49 6 313
Psittaciformes 78 16 358
Cuculiformes 11 2 143
Strigiformes 20 1 178?
Caprimulgiformes 11 ie) 105?
Apodiformes 39 3 103
Trogoniformes 3 10) 39
Coraciiformes 20 [e} 152
Piciformes 14 J 355
Passeriformes 499 50 5,712
REPTILES 169 38 4,771
Testudines 78 11
Rhynchocephalia 1 (0) 2?
1. Biological Diversity
Table 17.1 IUCN Threatened Vertebrates (1990 Red List)
CLASS NUMBER OF SPECIES APPROXIMATE TOTAL
ORDER THREATENED ENDANGERED OF DESCRIBED SPECIES
REPTILES (continued)
Sauria 43 9 2,000
Serpentes 33 7 2,500
Crocodylia 15 11
AMPHIBIANS 57 8 4,014
Caudata 25 2
Anura 32 6
FISHES 713 368 20,000
LAMPREYS 3 {e}
SHARKS, etc. 3 te)
BONY FISH 707 368
TOTAL VERTEBRATES 2,475 686 42,784
Sources: World species totals for groups of animals are derived from the following sources - mammals: Corbet, G.B. and Hill, J.E. 1991.A World
List of Mammalian Species. 3rd edn. Natural History Museum, London and Oxford Univerity Press, Oxford; birds: Sibley, C.G. and Monroe, B.
L. 1990. Distribution and Taxonomy of Birds of the World. Yale University Press, New Haven & London; reptiles, amphibians and fishes: various
sources.
Note: Table only includes groups of animals of which one or more species are listed as threatened, with the exception of mammals for which all
orders are included. Species categorised as Extinct are not included, those as Extinct? are.
Table 17.2 1UCN Threatened Invertebrates (1990 Red List)
NO. OF SPECIES APPROXIMATE TOTAL
PHYLUM CLASS THREATENED ENDANGERED OF DESCRIBED SPECIES
CILIOPHORA 1 0) ?
CNIDARIA 154 0) 9,000
PLATYHELMINTHES TURBELLARIA 4 2 12,700
NEMERTEA 10 ) 650
MOLLUSCA 409 85 50,000
ANNELIDA 139 2 8,700
ARTHROPODA INSECTA 1,083 56 750,000
MEROSTOMATA 4 ) 4
ARACHNIDA 18 1 68,000
CRUSTACEA 126 3 42,000
ONCHYOPHORA 27 ) 65
ECHINODERMATA 2 ) 6,000
TOTAL INVERTEBRATES 1,977 149 947,119
Sources: various.
Notes: Table only includes groups of animals of which one or more species are listed as threatened.
Species categorised as Extinct are not included, those as Extinct? are.
Threatened Species
Table 17.3 Country totals of threatened plants and vertebrates
PLANTS MAMMALS BIRDS REPTILES AMPHIBIANS FISH
ASIA 6608 497 918 146 9 124
Afghanistan 4 13 13 1 1 (9)
Bahrain 0 1 4 0 0 1
Bangladesh 33 15 27 14 i) 0
Bhutan 15 15 10 1 0 0
British Indian Ocean Territory _ (0) (e) (0) (°) ts) (0)
Brunei 40 9 10 3 ie) 2
Cambodia 11 21 13 6 (0) 5
China 350 40 83 7 1 7
Cyprus 43 1 17 1 0 0
HongKong 5 1 9 2 () ()
India 1336 39 72 ts 3 2
Indonesia 70 49 135 13 0 29
Iran, Islamic Rep 301 15 20 4 0 2
Iraq 1 9 17 0 0 2
Israel 3 8 15 1 1 ()
Japan 41 5 31 0 1 3
Jordan 752 5 11 0 0 0
Korea, Dem People’s Rep (0) 5 25 (0) 0 0
Korea, Rep 33 6 22 te} 0 0
Kuwait 1 5 7 (e) 0 te)
Laos 3 23 18 5 0 5
Lebanon : 5 4 15 1 0 ()
Malaysia 522 23 35 12 0 6
Maldives 1) 1 1 te) 0 (0)
Mongolia 0 9 13 (0) (0) 0
Myanmar 23 42 10 0 2
Nepal 33 22 20 9 (0) 0
Oman 2 6 8 te) tt) 2
Pakistan 14 15 25 6 0 0
Philippines 159 12 39 6 0 21
Qatar 0 0 3 0 i) 0
Saudi Arabia 2 9 12 (0) (°) te)
Singapore 19 4 5 1 0 1
Sri Lanka 220 7 8 3 (0) 12
Syria 11 4 15 1 0 C)
Taiwan 95 4 16 10} 0 0
Thailand 68 26 34 9 0 13
Turkey 1944 5 18 5 1 5
United Arab Emirates 0 4 7 te) te) 0
Viet Nam 338 28 34 8 1 4
Yemen 134 6 9 0 0 0
‘USSR’ 20 38 3 oO 5
EUROPE 2677 ** 66 396 16 15 48
Albania 76 2 14 1 0 1
Andorra (¢) te) 1 0 0 )
Austria 25 2 13 (0) 0 2
Belgium 9 2 13 te) 0 1
Bulgaria 88 3 15 1 0 3
Czechoslovakia 29 2 18 0 0 2
Denmark 7 1 16 (0) (°) te)
Faeroe Islands (0) (0) 2 (e) te) (°)
Finland 11 3 12 0 0) 1
France 143 6 21 2 il 3
Germany ae 2 17 0 0 3
Greece 526 4 19 3 te) 6
Hungary 21 2 16 e) 0 2
Iceland 2 1 2 0 0 1
lreland 4 0 10 0 0 1
Italy 210 3 19 2 7 3
Liechtenstein (0) te) 3 (0) 0 0
Luxembourg 1 1 8 ie) t¢) ts)
Malta 4 0 13 tt) 0 0
Monaco () (e) 0 te) 0 ()
Netherlands 7 2 13 te) 0 1
Norwa' 13 3 8 i¢) 0 1
Polan 16 4 16 0 0 1
Portugal 240 6 18 0 1 0
Romania 67 2 18 iJ (0) 4
San Marino (0) 0 0 0 0 0
Spain 936 6 23 5 3 2
Sweden 10 1 14 0 0 1
Switzerland 18 2 15 ie} 1 3
United Kingdom 24! 3 22 te} (o} 1
Vatican City 0 (0) ts) (0) 0 t)
Yugoslavia 190 3 17 1 2 5
NORTH AND CENTRAL AMERICA 5747 145 219 88 27 27T
Anguilla (0) 0 (0) 0 0
Antigua and Barbuda 1 ° 2 (0) 0 0
Aruba 0) 0 (0) 0 0 0
Bahamas 24 2 4 3 0 ()
Barbados 1 1 1 0 (°) tt)
Belize 36 8 4 3 0 0
Bermuda 1 (0) 2 te) () ts)
Canada 12 5 6 0 i) 15
Cayman Islands 0 te) 2 2 0 ts)
Costa Rica 419 10 14 2 0 0
239
1. Biological Diversity
Table 17.3 Country totals of threatened plants and vertebrates (continued)
PLANTS MAMMALS BIRDS REPTILES AMPHIBIANS FISH
NORTH AND CENTRAL AMERICA (continued)
Cuba 860 11 15 4 (0) te)
Dominica 62 t) 3 0 0 0
Dominican Republic 50 1 5 4 0 0
El Salvador 26 6 2 1 0 0
Greenland (Denmark) () 2 1 0 (t) 0
Grenada 4 t) 2 i) 0 0
Guadeloupe 14 t) 1 0 tt) tt)
Guatemala 282 10 10 4 0 0
Haiti 13 1 4 4 (0) ()
Honduras 43 7 1 3 () (*)
Jamaica 10 5 2 3 te) ()
Martinique 12 0 3 0 0 0
Mexico 883 25 35 16 4 98
Montserrat 1 0 1 0 0 0
Netherlands Antilles () () 3 2 () ()
Nicaragua 68 8 7 2 0 0
Panama 549 13 14 2 0 0
Puerto Rico 84 2 4 5 1 (°)
St Lucia 3 te) 5 (°) () (:)
St Vincent and the Grenadines 0 3 (*) 0 ()
St Kitts and Nevis 0 0 1 te) 0 0
Trinidad and Tobago 5 1 3 (°) (°) ()
Turks and Caicos Islands 1 ts) (0) 1 (1) ts)
United States 2262 27 43 25 22 164
Virgin Islands (British) 1 () 3 1 fe) ()
Virgin Islands (US) 10 0 3 1 0 ts)
SOUTH AMERICA 2061 239 535 58 2 14
Argentina 159 23 53 4 1 1
Bolivia 39 21 34 4 0 iu
Brazil 318 40 123 11 0 9
Chile 284 9 18 0 0 1
Colombia 327 25 69 10 te) ()
Ecuador{a} 256 21 64 8 te) 0
French Guiana 47 10 5 2 0 0
Guyana 68 12 9 3 0 1
Paraguay 15 14 34 4 0 0
Peru 360 29 75 6 1 1
Suriname 68 1 6 1 t) 0
Uruguay 14 5 11 2 0 0
Venezuela 106 19 34 3 0 t)
OCEANIA 2673 60 168 21 7 18
American Samoa 1 1 (0) 0 0
Australia 2024 38 39 9 3 16
Cook Islands ) (e) 1 te) te) (¢)
Fiji 25 2 1 5 4 1 ()
French Polynesia 65 (¢) 20 () (°) [}
Guam 12 2 4 0 0 0
Kiribati t°) te) 2 (0) te) ()
Marshall Islands (0) te) 1 0 0 ()
Micronesia, Federated States of 1) 5 3 1 0 0
Nauru 0 ie} 2 i) oO 0
New Caledonia 168 1 5 (0) Y) (¢)
New Zealand 232 1 26 1 3 2
Niue 0 0 0 (0) (0) (*)
North Marianas Islands 8 1 2 () 0 (*)
Palau i} 1 3 (¢) 0 (t)
Papua New Guinea 88 5 25 1 i) t°)
Pitcairn Island 3 te) 1 0 (*) ()
Solomon Islands 28 2 20 3 0 1)
Tokelau te) te) (0) te) t) tt)
Tonga c') i) 2 1 0 o
Tuvalu 0 0 1 ts) (*) t)
Vanuatu 8 1 3 1 0 0
Wallis and Futuna Islands 0 0 0 (*) tt)
Western Samoa 12 1 2 (°) t') tt)
ANTARCTICA 4 0 0 t) tt) Oo
Antarctica 0 0 ie) 0 0 ts)
Falkland Islands (Malvinas) 4 ) te) 0 0 0
French Southern Territories 1°} ie) ie) i) i) tt)
AFRICA 3308 688 453 89 8 49
Algeria 145 12 15 0 0 1
Angola 19 14 12 2 0 0
Benin 3 11 1 2 (°) tt)
Botswana 4 9 6 1 0 0
Burkina Faso (¢) 10 1 2 0 oO
Burundi ts) 4 5 1 tt) 0
Cameroon 74 27 17 2 1 11
Cape Verde 1 t) 3 1 te) t)
Cental African Rep te) 12 2 2 te) t)
Chad 14 18 4 2 oO 0
Comoros 3 3 5 0 0 1
Congo 4 12 3 2 te) t)
Cote d'Ivoire 70 18 9 1 1 0
Djibouti 3 6 3 () (0) te)
Egypt 91 9 16 2 0 1
240
Threatened Species
Table 17.3 Country totals of threatened plants and vertebrates (continued)
PLANTS MAMMALS BIRDS REPTILES AMPHIBIANS FISH
AFRICA (continued)
Equatorial Guinea 8 15 3 2 1 0
Ethiopia 44 25 14 1 0 0
Gabon 80 17 4 2 0 0
Gambia 0 i 1 2 0 0
Ghana 34 13 8 2 0 0
Guinea 36 17 6 1 1 (0)
Guinea-Bissau 0 5 2 2 0 0
Kenya 144 Ue 18 2 (:) tt)
Lesotho 7 2 7 0) 0 tt)
Liberia 1 18 10 2 i?) 0
Libya 58 12 9 1 0 0
Madagascar 194 50 28 10 0 0
Malawi 61 10 7 1 0 0
Mali 15 16 4 2 te) 0
Mauritania 3 14 5 1 () 0
Mauritius 269 3 10 6 0 0
Mayotte 0 1 (0) 0 0
Morocco 194 9 14 (0) 0 1
Mozambique, People’s Rep 89 10 1 1 0 1
Namibia 17 11 7 2 0 4
Niger 1 15 1 1 0 0
Nigeria 9 25 10 2 0 0
Reunion 96 te) 1 te) 0 ()
Rwanda 0 1 Uf 2 O i)
St Helena (0) 1 0 0 0
Sao Tome and Principe yo 1 7 0 0 0
Senegal 32 1 5 2 t¢) 0
Seychelles 75 1 9 2 3 0
Sierra Leone 12 13 7 2 0 0
Somalia 52 17 7 1 () tt)
South Africa 1016 25 13 3 1 28
Sudan 9 17 8 1 0 i)
Swaziland 25 tt) 5 1 () ()
Tanzana 158 30 26 3 0 0
Togo t) 9 1 2 i} i)
Tunisia 26 6 14 1 0 0
Uganda 1 16 12 1 0 0
Western Sahara 0 5 5 0 i*} i)
Zaire 3 31 27 2 te) 1
Zambia 1 10 10 2 ie} 0
Zimbabwe 96 9 6 1 0 0
Sources: IUCN 1990. 1990 IUCN Red List of Threatened Animals. IUCN, Gland and Cambridge. Additional range data from WCMC Animal
Database and other sources. Bird ranges from Sibley, C.G. and Monroe, B.L. 1990. Distribution and Taxonomy of Birds of the World. Yale
University Press, New Haven and London.
Notes: Plants: numbers include many taxa below species level and also Ex/E species. Vertebrates: marine species are excluded. Extinct taxa are
excluded, Only full species are accounted for. Includes K categories - i.e. all threatened species as defined by IUCN. Mammals: cetaceans are
excluded. Birds: the countries within the breeding and wintering range are included (where data available). Fishes: not included are c. 252 spp. of
Lake Victoria cichlids, many thought to be extinct or severley threatened. ** excludes figures for Germany (German Dem Rep = 11; Germany,
Fed Rep = 15). ' includes Gibralter (UK = 23; Gibralter = 1). ? includes Rotuma. ® total for Sao Tome only.
The majority of threatened mammalian species occur in
mainly tropical countries, with highest numbers recorded
from Madagascar (53), Indonesia (49), China (40) and
Brazil (40). India, Australia, Zaire and Tanzania also have
large numbers of species at risk, as do Mexico, USA,
‘USSR’ and most South American and Southeast Asian
countries.
A regression analysis (Fig. 17.4) shows that Madagascar
and Indonesia in particular have more threatened species in
relation to country area than would be predicted statistically
(points above the line) whereas USA, for example, has
fewer.
There are approximately twice as many threatened bird
species as mammals (1,029 versus 507) but they show a
similar distributional pattern. The majority are concentrated
in southern and Southeast Asia, USA, Mexico, and South
America. The ten countries listed all have more than 40
threatened species. In comparison, Europe, Africa, Canada,
the Middle East and the Arabian Peninsula have relatively
few globally threatened bird or mammal species.
241
Figure 17.3 Per cent of known species
classed as Endangered
Per cent endangered
Mamma | s
Fish
Birds
Reptiles
Amphibians
All Invertebrates
Taxonomic grouping
1. Biological Diversity
Table 17.4 Countries with greatest numbers of threatened vertebrates
MAMMALS BIRDS REPTILES AMPHIBIANS FISHES
COUNTRY TOTAL COUNTRY TOTAL COUNTRY TOTAL COUNTRY TOTAL COUNTRY TOTAL
Madagascar 53 Indonesia 135 USA 25 USA 22 USA 164
Indonesia 49 Brazil 123 India 17 Italy 7 Mexico 98
Brazil 40 China 83 Mexico 16 Mexico 4 Indonesia 29
China 40 India 72 Bangladesh 14 Australia 3 South Africa 28
India 39 Colombia 69 Indonesia 13 India 3 Philippines 21
Australia 38 Peru 65 Malaysia 12 New Zealand 3 Australia 16
Zaire 31 Ecuador 64 Brazil 11 Seychelles 3 Canada 15
Tanzania 30 Argentina 53 Colombia 10 Spain 3 Thailand 13
Peru 29 USA 43 Madagascar 10 Yugoslavia 2 Sri Lanka 12
Viet Nam 28 Myanmar 42 Myanmar 10 Cameroon 11
Sources: IUCN 1990. 1990 IUCN Red List of Threatened Animals. (UCN, Gland and Cambridge; WCMC 1991. The World Conservation
Monitoring Centre Animal Database. WCMC, Cambridge; Bird ranges estimated from Sibley, C.G. and Monroe, B.L. 1990. Distribution and
Taxonomy of Birds of the World. Yale University Press, New Haven and London.
Notes: Mammals - Cetacea (whales, dolphins) are excluded; Birds - estimates include breeding and overwintering species (where data available);
Reptiles - marine turtles are excluded; Fishes - the estimates do not include c. 252 species of cichlids in Lake Victoria. Marine species are also
excluded. Extinct taxa in all groups are excluded; only full species, not subspecies, are accounted for. Numerous countries had 1 threatened
amphibian species, therefore the last row could not be filled for this column.
Several factors may be involved in this distribution. Other
things being equal, the number of threatened species in a
country should be correlated with the total number of
species present, and tropical countries generally have a
higher species richness than temperate ones. The high
current rate of human population increase, and consequent
high rates of habitat loss and modification in tropical
countries, is doubtless an important factor.
The global distribution of species richness, the non-
matching and uneven geographic spread of conservation
activity and field survey work, and the patchy review to
which most taxonomic groups have been subjected jointly
mean that the IUCN Red List gives an as yet incomplete
picture of the global distribution of species which may be
under threat.
Habitat distribution of threatened animals
Information on habitat requirements is not consistently
available for all threatened species. A useful indication of
the global situation can be derived from analysis of the
threats facing, and habitat types occupied by, the mammals
of Australasia and the Americas and the birds of the world.
As stated above, habitat loss or modification is the main
category of threat affecting these species. The two habitat
types in which the largest number of threatened mammals
occur are lowland tropical rain forest (TRF) (37%) and
montane TRF (19%), which together are occupied by 43%
of all threatened Australian and American mammal species
(Fig. 17.5). Both these habitat types are found exclusively
in tropical regions, between latitudes 28°S and 28°N. Other
tropical and subtropical habitats such as dry savanna, humid
savanna, desert and semi-desert also possess large numbers
of threatened mammals. In contrast, temperate and polar
242
habitats such as coniferous and boreal forest, Mediterranean
forest and scrub, tundra and polar ice harbour relatively
few threatened species.
In general the world’s threatened bird species occupy a
range of habitat types remarkably similar to the threatened
mammals of Australasia and the Americas, with 43%
occurring in TRF. The percentages occurring in marine,
freshwater, grassland and polar habitats are also very
similar, but there are some notable differences (Fig. 17.6).
The major disparity is that some 38% of threatened birds
are found on oceanic islands. These are primarily flightless
or ground-nesting species which are threatened by
introduced predators, for example rats and mongooses. A
direct comparison with mammals is not possible because
Thornback and Jenkins did not include oceanic islands;
however, there are few mammals on such islands. A higher
percentage of threatened birds than mammals occurs in
seasonal woodlands (20% v. 8.4%), while this trend is
reversed in arid (1% v. 14%), and coastal and estuarine
habitats (5% v. 14%).
Madagascar has the highest number of threatened mammal
species (50). Most of these are forest-dwelling lemurs.
Harcourt and Thornback (1990) identify habitat destruction
as the main threat to lemurs, and estimate that at current
rates of cutting (1.2% per year) only forests on the steepest
slopes will survive the next 35 years.
AQUATIC HABITATS
These systems have received little attention in comparison
with terrestrial habitats, and very little survey work has so
far been undertaken in tropical areas. A recent synthesis
(Moyle and Leidy, in press) demonstrated that fishes
provide reliable indicators of trends in aquatic diversity.
Threatened Species
Figure 17.4 Relationship between number of threatened species and country area
Number of threatened mammals
Tanzania
cc [0 ete en en a nas! » See Sees ene = Pope pg nnn nnn n eee n anne nn neen eee e ewe nn enna wen en nnn nee n enn eenenreeeeenneneennnne=
United States
K
A OK
25 haitas 2K Me crrccetrecceccenceneeneenteeeeneeeneceeteneeceeeeteetneeeeceeacery
South Africa
*K
Argentina
20 Eee ee ee eee ee ee |
10, 000 20, 000 50,000 100, 000 200, 000 500, 000 1,000,000 2,000,000
Country area (1000 ha)
Figure 17.5 Habitat distribution of threatened mammals
40
§
Montane rain forest
Lowland rain forest
Humid savanna
Desert & semi-desert
Dry savanna
Alpine & montane
Coastal
Freshwater
_
fo)
Per cent of threatened mammals occupying habitat type
8
Coniferous & boreal forest
Habitat Type
Note: Data for Australasia and the Americas, excludes cetaceans.
243
1. Biological Diversity
Figure 17.6 Habitat distribution of threatened mammals and birds
50
Boo}
=
2
a
ce
aa
2
2
8
he PUM =
a
geal Mammals
Birds
Tropical Grassland Coasts & Arid Freshwater Mountains Seasonal Marine Tundra& Oceanic
forest heath& estuaries woodlands polar ice Islands
scrub
Habitat type
Note: Mammal data for Australasia and the Americas, excludes Cetacea; bird data are global.
Information on the fish faunas of North America, Europe,
Iran, South Africa, Sri Lanka, Australia, Costa Rica, Brazil
and Chile was analysed. The well-supported conclusion of
this review was that at least 20% (c. 1,800 species) of the
world’s freshwater fish species are seriously threatened or
extinct. Declines usually resulted from cumulative effects of
several long-term factors. Habitat modification (competition
for water, drainage, pollution), introduced species and
commercial exploitation were identified as the major causes
of decline. Recent fieldwork in Madagascar (Reinthal and
Stiassny, 1991) corroborates these general conclusions: the
native fish fauna in eastern and central Madagascar had
declined severely because of introductions and habitat
degradation as a result of forest clearance. These trends,
coupled with inadequate knowledge of freshwater faunas
and the strong representation of freshwater species in the
list of known extinct species (see Chapter 16), indicate that
aquatic systems require increased conservation attention.
THREATENED SPECIES ON ISLANDS: PLANTS
About one in six plant species grows on oceanic islands;
one in three of all known threatened plants are island
endemics. This is a measure of the diversity and fragility of
island ecosystems and their importance in _ plant
conservation.
Damage to most island floras occurred in the era of
European exploration and colonisation, when oceanic
islands became strategically important to the maritime
powers. Most island floras evolved in the absence of large
grazing animals and few endemic plants had defences
against grazing animals.
On St Helena, goats were introduced in 1513 and within 75
years had formed vast herds. Botanists only reached the
island in 1805-10, long after the damage had been done,
and so one can only speculate on the original flora. Today
46 endemic species are known, seven of them extinct
(Cronk in litt., 1991), but J.D. Hooker estimated that there
must have been originally over 100 endemic species (quoted
in Lucas and Synge, 1978). Most of these species will
never be known.
Philip Island, near the penal colony of Norfolk Island, has
been affected even more severely. The island was believed
to have carried a mixture of scrub and dense forest when
discovered by Captain Cook in 1744. The introduction of
goats, pigs and later rabbits reduced this vegetation to a
near desert in which by 1964 the endemic Philip Island
Glory Pea Streblorrhiza speciosa had become extinct and
the endemic hibiscus Hibiscus insularis reduced to four
aged bushes.
Whereas goats, sheep, pigs and even rabbits can be
controlled and even eliminated, the problem of introduced
plants is much more intractable. Enthusiastic gardeners
often brought to islands the plants they used to grow at
home, and some of these plants proved to be devastatingly
invasive in the native vegetation, outcompeting the native
flora. In Mauritius, for example, visitors today see rich
green thickets and forests covering the hills, but few realise
that virtually all this vegetation is of introduced plants. The
only viable strategy for saving the Mauritian endemic flora
in the short term is to make small weeded plots within the
forest, a few hectares at a time. Other islands where the
native flora is greatly threatened by introduced plants
include Rodrigues, St Helena, Hawaii and Juan Fernandez.
It is noticeable that introduced plants tend to be much more
destructive of island ecosystems than of continental ones.
Following the TDWG geographical classification (see
Chapter 14), there are about 80 islands or island groups
with significant endemic floras (here defined as more than
five endemic species). For nearly half of these islands, a
detailed assessment has been made of which species are
threatened (Table 17.6).
Degree of threat to species varies greatly from one island
or island group to another. Islands with severely affected
floras include:
e Hawaii: 108 endemic taxa have already gone extinct, 15
are either Extinct or Endangered, 138 are Endangered,
37 are Vulnerable, 126 are Rare, and 9 are
Indeterminate - a total of 433 threatened taxa. Hawaii
has, therefore, one of the most distinctive and one of the
most threatened floras in the world.
St Helena, in the Atlantic Ocean, where all of the 46
endemic known species are threatened, 7 of them Extinct
and 19 Endangered
Bermuda, north of the Caribbean: all but one of the 15
endemic species are threatened, 3 of them Extinct and 4
Endangered
Rodrigues, a dependency of Mauritius in the Indian
Ocean: all but 2 of the 45 endemic species are
threatened, 27 of them Endangered or Extinct
Norfolk Island, east of Australia: where all but 2 of the
36 endemic species are threatened, 1 of them Extinct and
11 Endangered.
On each of these islands, the native plants are reduced to
small patches of relict vegetation, and often have
populations of ten individuals or fewer. It is, however,
encouraging to see that on all the four islands listed above,
there are active programmes to rescue the threatened plants
although it may take centuries to restore the native
vegetation.
Other islands have fared better. For example, the native
forests on Lord Howe Island, a dependency of Australia,
are still intact and are now well protected in a national
park. Of the 84 endemic species in the Table, only one is
Extinct and three Endangered, but 72 are Rare, meaning
their world populations are low but they are not under
threat. Among coral islands, the important endemic floras
of Aldabra (Indian Ocean) and Henderson Island (Pacific
Ocean) are intact and both are now effectively protected as
nature reserves.
For some of the islands with larger floras, the flora has
only been partly assessed. The true numbers of threatened
species may be higher than those quoted. This is probably
true, for example, of Cuba and Jamaica, with their very
large endemic floras.
The islands listed in Table 17.5 all have more than 10
endemic species of plants but the conservation status of
those plants is not known. The immediate priority here is
for field surveys to assess the situation and provide a basis
for conservation action. For more details see Table 14.1.
245
Threatened Species
Table 17.5 ‘Priority islands for surveys
of endemic flora
ESTIMATED ENDEMIC
PLANT SPECIES
AFRICA
Annobon 17
Bioko 49
Cape Verde 92
Principe 35
Sao Tome 108
CARIBBEAN
Bahamas 112
Cayman Is 18
Dominican Republic/Haiti 1800
Trinidad-Tobago 215
Virgin Is, US and British 28 +
Most Lesser Antillean Islands total 327
INDIAN OCEAN
Andaman Is 144
Nicobar Is 72
Comoros 136
PACIFIC
American Samoa 27
Coco, Isla del 15
Fiji 700
Marquesas Is 105
New Caledonia 2480
Northern Marianas 81
Society Is ?
Taiwan 892
Tonga 25
Tuamotu Is 20
Tubuai Is 140
Vanuatu 150
Western Samoa 57
Table 17.6 covers higher plants (flowering plants, ferns,
gymnosperms) endemic to the island or island group
concerned. The main figures are of species; the figures in
brackets are of additional endemic infraspecies (subspecies
and varieties). Where an endemic species is divided into
several infraspecies in the database, it has been counted
only at the infraspecies level; however the parallel table in
Chapter 14 adds these endemic species into the endemic
species totals. Thus, the total for Mauritius here is 236 (54)
but above is 246 species, since the 54 infraspecies include
10 species that are wholly endemic to Mauritius.
THREATENED SPECIES ON ISLANDS: BIRDS
Islands are important for bird conservation: over 1,750
species (some 17% of the world’s bird species) are confined
to islands and of these, 402 (23%) are threatened (Johnson
and Stattersfield, 1990) compared with only 11% of birds
worldwide (Collar and Andrew, 1988). In addition, island
birds have suffered the majority of bird extinctions which
have occurred during historic times.
Distribution of island endemics
A high proportion of threatened island species are
concentrated in a few geopolitical units: a total of 92 such
units have one or more threatened species; 11 of these
(Cuba, Hawaiian Islands, Indonesia, Marquesas Islands,
Mauritius, New Zealand, Papua New Guinea, Philippines,
Sao Tome, and Principe, Seychelles and Solomons) support
iversi
1. Biological D
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246
over half the threatened species restricted to islands. Over
90% of threatened species restricted to islands are endemic
to their geopolitical units, with a few island groups having
particularly large numbers of threatened endemics (e.g.
Indonesia 91, and the Philippines 34). Some 25 islands
support a single threatened endemic only.
After Indonesia and the Philippines, the islands of the
Pacific Ocean support the largest number of threatened
species (110). Although when compared to the Atlantic
islands this constitutes a much lower portion of the
endemics occurring in the region (38% and 50%,
respectively) it nonetheless accounts for 27% of threatened
species restricted to islands.
Degree of threat
Of the 402 species restricted to islands, the greatest number
of those considered Endangered or Vulnerable occur within
the Pacific region: 31 of the 66 Endangered species and 29
of the 71 Vulnerable species. These include a wide range
of species, such as the severely endangered Barred-wing
Rail Nesoclopeus poeciloptera, known only from Fiji, and
the New Caledonian endemic Kagu Rhynochetos jubatus,
belonging to a monotypic family and therefore regarded as
a high priority for conservation action.
Habitat requirements
The majority of threatened island birds are forest species.
Rain forest supports 200 (50%) of the threatened species.
Lowland and montane forests contribute almost equally,
being used by 101 and 112 species, respectively (42 species
use both types; 29 rain forest species could not be assigned
to the lowland/montane division). The other major forest-
type, seasonal/temperate forest, supports 113 species. In
total, forests of all categories support 310 species,
accounting for 77% of threatened island endemics.
Threats
The most important factor threatening island species is
habitat destruction, affecting over 50% of threatened island
247
Threatened Species
species. Given the number of extinctions attributable to
introductions, it is of interest that introduced species now
appear to be a major threat to only 20% of threatened island
endemics, a much smaller proportion than might be
expected and a considerably smaller proportion than the
41% of island species which are at risk simply by having a
limited range. Other factors (hunting, trade, human
disturbance, natural causes and fisheries) each affect less
than 10% of threatened island birds. For some 60
threatened island endemic birds, further field research is
needed to identify the cause of decline.
References
Collar, N.J. and Andrew, P. 1988. Birds to Watch: the ICBP world
checklist of threatened birds. ICBP Technical Publication No.8,
ICBP Cambridge, UK.
Corbet, G.B. and Hill, J.E. 1991. A World List of Mammalian Species.
Third edition. Oxford University Press, UK.
Diamond, A.W. 1987. Save the Birds. ICBP, Girton, Cambridge.
Harcourt, C. and Thornback, J. 1990. Lemurs of Madagascar and the
Comoros. The IUCN Red Data Book. TUCN, Gland, Switzerland
and Cambridge, UK.
IUCN, 1990. 1990 IUCN Red List of Threatened Animals. TUCN,
Gland and Cambridge.
Johnson, T.H. and Stattersfield, A.J. 1990. Global review of island
endemic birds. [bis 132:167-180.
Lucas, G. and Synge, H. 1978. The IUCN Plant Red Data Book.
IUCN, Switzerland.
Mace, G.M. and Lande, R. 1991. Assessing extinction threats: toward
a re-evaluation of JIUCN threatened species categories.
Conservation Biology 5(2):148-157.
Moyle, P.B. and Leidy, R.A. (in press). Loss of biodiversity in
aquatic ecosystems: evidence from fish faunas. In: Feidler, P.L.
and Jain, S.K. (Eds), Conservation Biology: the theory and practice
of nature conservation, preservation, and management. Chapman
and Hall, New York.
Reinthal, P.N. and Stiassny, M.L.J. 1991. The freshwater fishes of
Madagascar: a study of an endangered fauna with recommendations
for a conservation strategy. Conservation Biology 5(2):231-243.
Sibley, C.G. and Monroe, B.L. 1990. Distributionand Taxonomy of
Birds of the World Yale University Press, New Haven and London.
Thornback, J. and Jenkins, M. 1982. The IUCN Mammal Red Data
Book Part 1. YUCN, Gland, Switzerland and Cambridge, UK.
The section on threatened plants on oceanic islands was
prepared by Hugh Synge.
1. Biological Diversity
18. GLOBAL HABITAT CLASSIFICATION
The world encompasses an enormous range of terrestrial
and aquatic environments, from polar ice-caps to forests,
and coral reefs to deep ocean trenches. The classification of
this immense range of variation into a manageable system
is a major problem in biology and underpins much of the
sciences of ecology and biogeography. It has not merely
theoretical interest, but is of fundamental importance in the
management and conservation of the biosphere.
Within ecology, a wide variety of terms has been coined -
community, habitat, ecosystem, biome - intended to help in
such a classification. Some of these can be seen as forming
a loose and ill-defined hierarchy analogous in some ways
with the taxonomic system developed for classifying
organisms, discussed fully in Chapters 2 and 3. However,
the classification of the natural environment is far more
problematic than the classification of organisms and none of
the above terms has a rigid, satisfactory and universally
accepted definition. Indeed there are good theoretical
grounds for questioning the basis of such a classification.
This is because these systems are ultimately based on an
assumption that the natural environment can be divided into
a series of discrete, discontinuous units rather than
representing different parts of a highly variable natural
continuum, whereas in reality the latter is undoubtedly a
more accurate description of the world.
In general, attempts to classify ecological units are based on
identification of the species which occur in them along with
a description of the physical characteristics of the area.
Most terrestrial ecosystems, for example, are generally
identified on the basis of plant communities, that is areas
with similar plant species composition and structure. The
basic principle underlying this is that different species may
habitually be closely associated with each other over a wide
geographical range. The extent to which this is true is still
controversial - it can reasonably be argued that the
distribution of plant species is generally dependant on the
physical environment and historical accident rather than on
the occurrence or otherwise of other plant species, although
within a particular geographical region, species with similar
ecological requirements may, of course, be expected to
have similar distributions. Even if the concept of a
community is accepted, it is widely acknowledged that the
more rigidly a community is defined the more site-specific
it becomes and hence the more limited its use in analysis
and planning.
At the other extreme, very general habitat classifications
(‘forests’, ‘grasslands’, ‘wetlands’) are based on the
physical characteristics and appearance of an area,
independent of species composition. They cover such a
wide range of possible conditions that they have little
heuristic use: the term ‘forest’ applies both to highly
diverse lowland tropical rainforest and coniferous
monoculture, two systems which may have no, or virtually
no, species in common. Furthermore these general terms
are virtually impossible to define and delimit in a
universally applicable way. Thus, for example, the density
of tree cover necessary before an area can be called a
woodland is undefinable and any limit used will always be
arbitrary. Similarly, it is impossible to determine for how
248
long and how intensely an area must be flooded before it
can be classified as a wetland rather than a terrestrial
ecosystem. This naturally makes any mapping of habitats a
problematic task.
ECOSYSTEM MAPPING
Most global habitat classification systems have attempted to
steer a middle course between the complexities of
community ecology and the oversimplified terms discussed
above, although they too have the same problems of
definition and delimitation. Generally these systems will use
a more or less elaborate combination of a general definition
of habitat type with a climatic descriptor (e.g. ‘tropical
moist forest’, ‘temperate grassland’, ‘warm deserts and
semi-deserts’). Some systems also incorporate global
biogeography to take into account the floristic and faunistic
differences between regions of the world which may have
very similar climate and physical characteristics.
Further, ecosystem mapping may either take into account
man’s activities to attempt to produce a realistic,
contemporary map of land-cover types, or may create a
potential vegetation map from an analysis of climatic or
other environmental variables. The potential vegetation
maps produced from this approach are independent of actual
disturbances on the landscape.
Four of the major global classification systems are
presented here. The Classification of Biogeographical
Biomes of the World map (Plate 1), provides a modest
classification based largely on geography and potential
vegetation. The Ecoregions of the Continents map (Plate 2)
and the Major World Ecosystems map (Plate 3) are
produced from a combination of potential vegetation and
actual land-use. The Holdridge Life Zone Classification
map (Plate 5) depicts potential vegetation using the life zone
classification system developed by Holdridge (1967).
The map in Plate 1 depicts the terrestrial biogeographic
realms of the world and was produced for IUCN (Udvardy,
1975). This map provides a generalised framework to
represent the distribution of biogeographical regions, biotic
provinces, and biomes. The approach used to produce this
map utilised vegetation and forest maps to produce the map
categories. Over the past decade, this map has served IUCN
and UNESCO as a primary global biogeographical guideline
for conservation planning purposes. The distribution of
protected areas throughout the globe within these
biogeographical provinces is presented in Part 3.
The Ecoregions of the Continents Map in Plate 2 shows the
distribution of ecosystems at the regional scale across the
globe based upon existing climatic and vegetation data
Gailey, 1989a, b). The three levels of hierarchy used for
representing ecosystems on this map are domains, divisions,
and provinces. These categories are obtained by defining
aggregates of ecosystems into larger biome categories. This
map therefore represents a generalised depiction of
ecosystem distribution across the globe. Table 18.1 presents
the area contained in each region and its percentage of the
global land area.
Global Habitat Classification
Table 18.1 Ecoregions of the continents (Bailey)
ECOREGION DOMAINS, DIVISIONS, AND PROVINCES km2 Forcent
100 POLAR DOMAIN 38,038,000 26.00%
110 Icecap Division 12,823,000 8.77%
M110 Icecap Regime Mountains 1,346,000 0.92%
120 Tundra Division 4,123,000 2.82%
121 Polar deserts 283,000 0.19%
122 Arctic tundras 1,231,000 0.84%
123 Oceanic moss —and-—grass tundra 184,000 0.13%
124 Continental moss—and-lichen (typical) tundra 1,981,000 1.35%
125 Continental bush—and—shrub tundra 445,000 0.30%
M120 Tundra Regime Mountains 1,675,000 1.14%
M120 Tundra regime mountains (Antarctica) 60,000 0.04%
M121 Tundra—polar desert 795,000 0.54%
M122 Polar desert 820,000 0.56%
130 Subarctic Division 12,259,000 8.38%
131 Continental dark evergreen needleleaf open forest 2,285,000 1.56%
132 Continental light deciduous needleleaf open forest 1,286,000 0.88%
133 Eastern oceanic tayga 918,000 0.63%
134 Moderate continental dark evergreen needleleaf tayga 2,692,000 1.84%
135 Continental dark evergreen needleleaf tayga 1,880,000 1.29%
136 Continental and extreme continental light deciduous tayga 2,237,000 1.53%
137 Moderate continental small—leafed forest 251,000 0.17%
138 Continental mixed coniferous and small—leafed forest 710,000 0.49%
M130 Subarctic Regime Mountains 5,812,000 3.97%
M131 Open woodland—tundra 1,750,000 1.20%
M132 Open woodiand—creeping tees—tundra 1,806,000 1.23%
M133 Forest—tundra of moderately and continental climate 686,000 0.47%
M134 Forest—creeping trees —tundra of extreme continental climate 1,203,000 0.82%
M135 Oceanic forest—tundra 367,000 0.25%
200 HUMID TEMPERATE DOMAIN 22,455,000 15.35%
210 Warm Continental Division 2,187,000 1.49%
211 Eastern oceanic mixed monsoon forest 65,000 0.04%
212 Moderate continental mixed forests 2,122,000 1.45%
M210 Warm Continental Regime Mountains 1,135,000 0.78%
M211 Oceanic forest—tundra 67,000 0.05%
M212 Oceanic forest—creeping tees 331,000 0.23%
M213 Forest—tundra of moderately continental and continental climate 736,000 0.50%
220 Hot Continental Division 1,670,000 1.14%
221 Permanently humid eastern oceanic broadleaf forests 788,000 0.54%
222 Moderately humid broadleaf forest in moderately continental climate 882,000 0.60%
M220 Hot Continental Regime Mountains 485,000 0.33%
M221 Forest—alpine meadows 485,000 0.33%
230 Subtropical Division 3,568,000 2.44%
231 Oceanic mixed constantly humid forests 3,568,000 2.44%
M230 Subtropical Regime Mountains 1,543,000 1.05%
M231 Forest—meadow of eastern oceanic (monsoon climate) 1,264,000 0.86%
M232 Oceanic constantly humid forest—alpine meadows 278,000 0.19%
240 Marine Division 1,347,000 0.92%
241 Oceanic meadow 92,000 0.06%
242 Western oceanic coniferous and mixed forests 210,000 0.14%
243 Permanently humid western oceanic broadleaf forests 951,000 0.65%
244 Western oceanic tayga 95,000 0.07%
M240 Marine Regime Mountains 2,194,000 1.50%
M241 Oceanic meadow-tundra 21,000 0.01%
M242 Oceanic forest—tundra 1,068,000 0.73%
M243 Forest—alpine meadows 1,105,000 0.76%
250 Prairie Division 4,419,000 3.02%
251 Temperate prairies (humid steppes and wooded steppes) of eastern parts 752,000 0.51%
of continents
252 Broadleaf—wooded steppes and meadow steppes of moderately 1,172,000 0.80%
continental climate
253 Small—leafed and coniferous wooded steppes of continental climate 787,000 0.54%
254 Open woodland, savannas, and shrub of eastern parts of continents 925,000 0.63%
255 Subtropical prairies (humid steppes and wooded steppes) of eastern 783,000 0.54%
parts of continents 0
M250 Prairie Regime Mountains 1,256,000 0.86%
M251 Continental steppe—forest—tundra and steppe—forest-meadow 690,000 0.47%
M252 Forest—alpine meadows 566,000 0.39%
260 Mediterranean Division 1,090,000 0.75%
261 Western oceanic mixed sclerophy/l forests and shrub 927,000 0.63%
262 Dry steppes and shrub of moderate continental climate 163,000 0.11%
M260 Mediterranean Regime Mountains 1,561,000 1.07%
M261 Forest—alpine meadows of western oceanic (mediterranean) climate 567,000 0.39%
M262 Shrub—forest—meadow of mediterranean climate 995,000 0.68%
300 DRY DOMAIN 46,806,000 32.00%
310 Tropical/subtropical Steppe Division 9,838,000 6.73%
311 Steppes and shrub of moderate continental climate 364,000 0.25%
312 Dry steppes, open woodland, and shrub of continental climate 846,000 0.58%
313 Shrub and semi—shrub semi-—deserts of continental climate 1,392,000 0.95%
314 Desert—like savannas, open woodland, and shrub 5,807,000 3.97%
315 Dry steppes and shrub of moderate continental climate 1,429,000 0.98%
M310 Tropical/subtropical Steppe Regime Mountains 4,555,000 3.11%
M312 Forest—meadow-—steppe of continental climate 670,000 0.46%
M313 Open woodland-—steppe of continental climate 2,714,000 1.86%
M314 Open woodland—shrub—desert 770,000 0.53%
M315 Open woodland-—steppe 400,000 0.27%
249
1. Biological Diversity
Table 18.1
ECOREGION DOMAINS, DIVISIONS, AND PROVINCES
300 DRY DOMAIN (continued)
320 Tropical/subtropical Desert Division
321 Shrub and semi—shrub semi—deserts and deserts of continental climate
322 Semi-—deserts and deserts
323 Inner continental shrub semi—desert
324 Inner continental deserts of continental climate
325 Western oceanic semi—deserts and deserts with high relative humidity
326 Inner continental semi—deserts and deserts of extreme continental climate
M320 Tropical/subtropical Desert Regime Mountains
M321 Desert—steppe and desert—steppe—desert of continental climate
M322 Extreme continental desert
M323 Desert—steppe
M324 Desert
330 Temperate Steppe Division
331 Dry steppes of continental climate
332 Steppes of moderately continental climate
333 Dry steppes of extreme continental climate
M330 Temperate Steppe Regime Mountains
M331 Forest—alpine meadows
M332 Continental open woodland—steppe
340 Temperate Desert Division
341 Semi—deserts and deserts of continental climate
342 Semi-—deserts of continental climate
343 Deserts of continental climate
344 Semi-—deserts of extreme continental climate
345 Deserts of extreme continental climate
M340 Temperate Desert Regime Mountains
M341 Extreme continental desert—steppe
400 HUMID TROPICAL DOMAIN
410 Savanna Division
411 Seasonally humid mixed (deciduous and evergreen) forests
412 Savannas, open woodland and shrub with seasonal moisture supply
413 Seasonally humid, predominantly deciduous forests
414 Humid tall—grass savannas and savanna forests
415 Moderately humid grassy savannas
416 Dry savannas and open woodland
M410 Savanna Regime Division
M411 Forest—steppe and forest—meadow of seasonally humid type
M412 Forest—-meadow, seasonally humid
M413 Forest—steppe, inner continental and leeward slopes
420 Rainforest Division
421 Eastern oceanic constantly humid forests
422 Mixed forests with short dry season
423 Constantly humid evergreen forests
424 Humid forests with short dry season
M420 Rainforest Regime Mountains
M421 Forest—meadow of constantly humid eastern oceanic type
M422 Forest—paramo and forest—meadow of constantly humid oceanic (and
windward—slope) type
M423 Forest—paramo and forest—-meadow
Ecoregions of the continents (Bailey)
km Percent
17,267,000 11.80%
1,321,000 0.90%
665,000 0.45%
3,674,000 2.51%
7,921,000 5.42%
958,000 0.65%
2,727,000 1.86%
3,199,000 2.19%
1,193,000 0.82%
899,000 0.61%
471,000 0.32%
636,000 0.44%
4,780,000 3.27%
1,790,000 1.22%
1,581,000 1.08%
1,409,000 0.96%
1,066,000 0.73%
893,000 0.61%
173,000 0.12%
5,488,000 3.75%
922,000 0.63%
1,213,000 0.83%
1,647,000 1.13%
399,000 0.27%
1,306,000 0.89%
613,000 0.42%
613,000 0.42%
38,973,000 26.64%
20,641,000 14.11%
1,346,000 0.92%
2,496,000 1.71%
4,951,000 3.38%
3,699,000 2.53%
4,771,000 3.26%
3,379,000 2.31%
4,488,000 3.07%
1,102,000 0.75%
1,220,000 0.83%
2,167,000 1.48%
10,403,000 7.11%
1,843,000 1.26%
2,893,000 1.98%
4,280,000 2.93%
1,387,000 0.95%
3,440,000 2.35%
728,000 0.50%
1,013,000 0.69%
1,700,000 1.16%
Source: Bailey, R.G. 1989. Ecoregions of the Continents. U.S. Department of Agriculture, Forest Service, Washington, D.C.
The map in Plate 3 (Olson et al., 1983) is a global ecology
map and represents the primary world ecosystem types as
of 1980. The regions on this map represent large areas
within which local ecosystems are present more or less in
a predictable fashion. Table 18.2 shows a country-by-
country breakdown of major ecosystem types along with an
estimate of the mass of carbon per unit area in live
vegetation in each country.
The Holdridge Life Zone Classification system, represented
in Plate 4, is a predictive scheme for identifying
undisturbed vegetation based generally upon the effects of
temperature, rainfall and evapotranspiration (Holdridge,
1967). This system was used to produce the map in Plate 5
that depicts the generalised distribution of eco-climatic
zones across the globe. Plate 5 therefore represents the
potential distribution of ecosystems in the world based on
a consideration of current climatic patterns. The Holdridge
Life Zone Classification System is described in further
detail in the Chapter 19.
ESTIMATING RATES OF CHANGE OF
ECOSYSTEMS
Given the difficulties of ecosystem definition and
delimitation outlined above, it is, unsurprisingly, extremely
difficult to measure existing areas of any given ecosystem
or habitat, and even more problematic to estimate rates of
loss. In large part this is because habitat alteration covers
a wide spectrum of change, from short-term, slight and
reversible disturbance to complete, and effectively
irreversible, destruction. Just as it is impossible to define
rigidly the limits of any given ecosystem or habitat, so it is
impossible to determine how much a given area of
ecosystem or habitat has to change before it can be
considered destroyed or converted. This problem is
compounded by the fact that the natural environment is not
static but rather dynamic, sometimes highly so, on a time
scale ranging from hours to millions of years. It is thus
difficult even to define an undisturbed ecosystem or habitat
as a standard against which to measure degree of
disturbance.
Global Habitat Classification
Table 18.2 Estimates of vegetation type and percent cover
OTHER DESERT & POLAR GRASS CROP & INTER—
COASTAL MAJOR SEMI-— AND AND SETTLE-— RUPTED MAJOR CARBON
AQUATIC WETLANDS DESERT ALPINE SHRUB MENTS WOODS FORESTS (Kg/m?)
WORLD 4% 2% 13% 12% 20% 11% 17% 22% 3.1
ASIA 4% 1%. 16% 9% 24% 17% 10% 18% 2.6
Afghanistan 11% 16% 62% 7% 4% We
Bangladesh 7% 5% 42% 25% 20% 46
Bhutan 25% 25% 19% 31% 3.7 *
Brunei 67% 33% 2.3 **
Cambodia 7% 4% 19% 5% 65% 6.5
China 1% 1% 14% 22% 21% 17% 5% 18% 2.4
Cyprus 100% 0.8 **
India 3% 0% 2% 2% 12% 44% 23% 14% ra
Indonesia 24% 9% 4% 9% 14% 40% 5.4
Iran, Islamic Rep 1% 30% 41% 7% 17% 4% 1.4
Iraq 30% 34% 33% 2% 1.0
Israel 40% 30% 30% War? ©!
Japan 21% 1% 5% 18% 27% 30% 4.2
Jordan 44% 49% 7% 0.6
Korea, Dem People’s Rep 12% 18% 14% 16% 39% 3.8
Korea, Rep 10% 24% 20% 24% 22% 3.4
Kuwait 9% 91% 0.3 *
Laos 7% 7% 86% 91
Lebanon 25% 75% 2'5)=*
Malaysia 8% 3% 2% 10% 33% 45% 6.7
Mongolia 32% 5% 52% 1% 3% 7% 1.4
Myanmar 5% 1% 2% 16% 23% 52% 6.7
Nepal 23% 35% 42% 44
Oman 13% 46% 34% 7% 0.5
Pakistan 1% 2% 21% 7% 46% 13% 9% 0% 11
Philippines 32% 2% 22% 13% 31% 47
Qatar 100% Ogu *
Saudi Arabia 1% 62% 33% 2% 1% 1% 0.5
Sri Lanka 32% 42% 3% 13% 10% 17
Syrian Arab Rep 18% 41% 36% 3% 1% 1.0
Taiwan 29% 12% 6% 53% 5.5 *
Thailand 7% 41% 7% 45% 5.5
Turkey 3% 9% 37% 18% 25% 7% 1.9
United Arab Emirates 3% 76% 21% 0.2
Viet Nam 17% 1% 4% 30% 49% 6.0
Yemen 7% 11% 74% 3% 6% 1.6
USSR (former) 3% 2% 5% 26% 10% 8% 21% 26% 3.2
EUROPE 6% 0% 9% 4% 35% 22% 23% 3.0
Albania 73% 27% 45 *
Austria 6% 36% 36% 22% 44
Belgium 40% 60% 21 *
Bulgaria 2% 59% 25% 14% 2.6
Czechoslovakia 29% 35% 36% 5.6
Denmark 31% 50% 19% 1.0 *
Finland 1% 12% 3% 12% 72% 5.3
France 5% 0% 2% 49% 28% 16% 2.9
Germany 3% 50% 24% 23% 3.8
Greece 26% 3% 40% 27% 3% 1.4
Hungary 82% 16% 2% 15
Iceland 9% 78% 13% 0.5
Ireland 3% 97% 0.8
Ital 19% 1% 35% 39% 6% 2.2
Luxembourg 100% 0.8 **
Netherlands 11% 84% 5% 0.8 *
Norwa' 6% 67% 1% 3% 5% 19% 2.2
Polan: 2% 4% 45% 38% 10% 28
Portugal 3% 12% 18% 45% 21% 3.6
Romania 3% 4% 54% 6% 34% 3.3
Spain 5% 0% 2% 16% 30% 25% 22% 3.2
Sweden 5% 22% 5% 13% 55% 44
Switzerland 24% 29% 29% 19% 3.6
United Kingdom 22% 27% 50% 1% 0.9
Yugoslavia 1% 56% 29% 14% 2.8
NORTH & CENTRAL AMERICA 5% 2% 3% 33% 9% 10% 17% 21% SA
Bahamas 40% 40% 20% OH!
Belize 17% 83% 10.8 *
Canada 5% 3% 44% 1% 3% 17% 27% 3.5
Costa Rica 33% 17% 17% 33% 3.5 *
Cuba 17% 5% 29% 29% 10% 12% 2.2
Dominican Rep 53% 26% 21% 0.7 *
El Salvador 20% 50% 30% eee
Greenland 1% 99% 0.5
Guatemala 12% 24% 65% 6.7
Haiti 21% 21% 57% Ova
Honduras 14% 22% 65% 7.9
Jamaica 67% 33% 1.0 *
Mexico 6% 0% 15% 0% 28% 13% 11% 27% 3.1
Nicaragua 2% 2% 24% 72% 8.6
Panama 17% 10% 27% 10% 37% 5:2
United States 5% 1% 5% 15% 16% 18% 24% 18% 3.1
251
1. Biological Diversity
Table 18.2 Estimates of vegetation type and percent cover
OTHER DESERT & POLAR GRASS CROP & INTER—
COASTAL MAJOR SEMI-— AND AND SETTLE— RUPTED MAJOR CARBON
AQUATIC WETLANDS DESERT ALPINE SHRUB MENTS WOODS FORESTS (Kg/m?4
SOUTH AMERICA 2% 3% 5% 2% 32% 8% 14% 33% 48
Argentina 3% 6% 26% 2% 23% 19% 17% 4% 15
Bolivia 0% 4% 1% 4% 50% 2% 27% 13% 3.3
Brazil 2% 2% 36% 6% 12% 42% 5.8
Chile 5% 21% 7% 20% 10% 7% 28% 24
Colombia 1% 3% 1% 29% 3% 13% 49% 6.8
Ecuador 5% 2% 16% 7% 16% 53% 6.1
French Guiana 4% 11% 7% 78% 9.6
Guyana 5% 4% 23% 4% 1% 63% 72
Paraguay 4% 10% 43% 43% 6.0
Peru 3% 5% 8% 18% 6% 19% 41% 5.5
Suriname 8% 33% 6% 53% 6.9
Uruguay 5% 3% 66% 26% 0.9
Venezuela 3% 5% 1% 49% 7% 9% 27% 4.2
OCEANIA 4% 1% 18% 0% 18% 5% 38% 16% 28
Australia 2% 20% 0% 19% 5% 40% 13% 2.6
Fiji 67% 11% 22% aan
New Caledonia 56% 11% 33% ie
New Zealand 21% 2% 25% 15% 21% 16% 27
Papua New Guinea 17% 12% 1% 2% 11% 57% 6.9
AFRICA 2% 2% 30% 0% 28% 7% 14% 17% 27
Algeria 0% 81% 4% 3% 8% 3% 07
Angola 0% 0% 1% 30% 4% 18% 46% 46
Benin 6% 76% 6% 12% 2.3
Botswana 4% 66% 5% 22% 3% 141
Burkina Faso 93% 3% 4% 25
Burundi 27% 36% 36% 3.0 *
Cameroon 4% 31% 7% 21% 37% 5.8
Cental African Rep 59% 5% 36% 5.0
Chad 1% 0% 49% 40% 1% 7% 1% 1.3
Congo 7% 15% 6% 17% 54% 7.6
Cote d'Ivoire 3% 37% 8% 12% 39% 5.7
Djibouti 11% 78% 11% 0.5 *
Egypt 3% 76% 8% 13% 0.5
Equatorial Guinea 100% 12.0 *
Ethiopia 0% 1% 2% 41% 4% 32% 20% 2.6
Gabon 7% 1% 9% 83% 10.4
Gambia 25% 50% 25% 2.8 *
Ghana 5% 1% 55% 2% 12% 25% 45
Guinea 1% 3% 77% 3% 1% 15% 2.8
Guinea-Bissau 25% 42% 33% ii7d ane
Kenya 3% 4% 35% 11% 42% 5% 25
Lesotho 75% 25% 1.4*
Liberia 11% 18% 18% 53% 7.3
Libya 1% 86% 9% 2% 2% 0.4
Madagascar 11% 45% 25% 15% 5% 1.8
Malawi 17% 2% 2% 29% 2% 46% 3.2
Mali 6% 40% 28% 4% 21% 1% 1.6
Mauritania 69% 16% 2% 13% 08
Morocco 8% 23% 1% 14% 16% 34% 4% 1.8
Mozambique, People’s Rep _ 6% 6% 18% 16% 17% 38% 4.0
Namibia 0% 1% 44% 33% 21% 1% 1.2
Niger 59% 26% 15% 0% 1.0
Nigeria 2% 2% 46% 24% 14% 12% 2.6
Rwanda 13% 13% 63% 13% ie
Senegal 2% 58% 38% 2% 2.2
Sierra Leone 12% 41% 12% 35% 47*
Somalia 11% 38% 36% 1% 14% 14
South Africa 4% 0% 15% 29% 13% 37% 2% 18
Sudan 1% 2% 24% 52% 10% 8% 2% 15
Swaziland 25% 75% 3.0 **
Tanzana 7% 31% 14% 17% 30% 3.4
Togo 10% 40% 10% 30% 10% 28
Tunisia 13% 23% 32% 15% 15% 1% 11
Uganda 15% 48% 15% 14% 9% 25
Western Sahara 2% 98% 0.3
Zaire 1% 8% 19% 3% 6% 63% 7.6
Zambia 1% 2% 7% 17% 73% 5.0
Zimbabwe 14% 18% 38% 31% 3.4
Source: Olson, J.S., Watts, J.A. and Allison, LJ. 1983. Carbon in Live Vegetation of Major World Ecosystems. Oak Ridge National Laboratory, Oak Ridge.
Olson, J.S., Watts, J.A. and Allison, LJ. 1985. Major World Ecosystem Complexes Ranked by Carbon in Live Vegetation: a database. Oak Ridge National
Laboratory, Oak Ridge.
Notes: For smaller countries the precision is limited by the 0.5 degree resolution of the Olson dataset. * entire country includes less than 20 42—degree cells;
** less than 5 cells
These problems notwithstanding, it is indisputable that man
is having an increasing effect on the natural environment
and that this effect extends to all the ecosystems considered
here. Some, very general, impression of this can be gained
252
from Table 18.3 which gives an estimate of the increase in
area of cropland between the years 1700 and 1980. It also
shows the decrease in forests and woodlands and grasslands
and pastures over that time. It is clear from these estimates
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that it is the forests and woodlands that have suffered the
most from conversion to croplands. Overall figures such as
these may mask other changes which are deleterious to
biological diversity. In Europe, for example, forest area has
actually increased during the twentieth century but this is
the result of large-scale planting of species-poor coniferous
monoculture; the area of species-rich natural and semi-
Table 18.3
VEGETATION TYPES
Global land use 1700-1980
AREA (10* km?)
1700 1850 1920
Forests and woodlands 6215 5965 5678
Grasslands and pasture 6860 6837 6748
Croplands 265 537 913
Global Habitat Classification
natural woodland has continued to decrease. Similarly, the
area of grassland in Europe has remained static or nearly so
over this period, but there has been wholesale conversion
from low nutrient-input, species-rich grassland, to high
input, intensively cultivated, species-poor pasture. It is
extremely difficult to map these changes and to measure
their effect.
CHANGE
PERCENTAGE AREA (mkm?)
1950 1980 1700-1980
5389 5053 -18.7% 11.62mkm?
6780 6788 -1.0% 0.72mkm?
1170 1501 +466.4% 12.36mkm?
Source: Richards, J.F. 1990. Land transformation. In: Turner, B.L. (Ed), The Earth as Transformed by Human Action.
Chapters 20 to 24 discuss five major habitat types (tropical
Tain forests, grasslands, wetlands, coral reefs and
mangroves) and assess the impact of man on each of these.
For only the first of these are adequate data available to
enable estimates for global rates of loss or conversion.
These are discussed at length, with individual case studies
adding detail. For the other systems considered, examples
of threats and changes are given, as well as indications of
their often considerable value to man.
References
Bailey, R.G. 1989a. Ecoregions of the Continents. U.S. Department of
Agriculture, Forest Service, Washington D.C.
253
Bailey, R.G. 1989b. Explanatory supplement to ecoregions map of the
continents. Environmental Conservation 16(4):307-309.
Holdridge, L.R. 1967. Life Zone Ecology. Tropical Science Center,
San José. 206pp.
Olson, J.S., Watts, J.A. and Allison, L.J. 1983. Carbon in Live
Vegetation of Major World Ecosystems. Oak Ridge National
Laboratory, Oak Ridge.
Olson, J.S., Watts, J.A. and Allison, L.J. 1985. Major World
Ecosystem Complexes Ranked by Carbon in Live Vegetation: a
database. Oak Ridge National Laboratory, Oak Ridge.
Richards, J.F. 1990. Land transformation. In: Turner, B.L. (Ed), The
Earth as Transformed by Human Action. Pp.163-178.
Udvardy, M.D.F. 1975. A Classification of the Biogeographical
Provinces of the World. \UCN Occasional Paper No. 18. IUCN,
Gland.
Part 1. Biological Diversity
19. BIODIVERSITY AND GLOBAL CLIMATE CHANGE
The currently increasing levels of the so-called ‘greenhouse’
gasses (e.g. carbon dioxide, methane, chlorofluorocarbons),
in the atmosphere could have large impacts on global
biochemical cycles and the climate system. This increase
results primarily from human industrial and agricultural
activities. There is currently a growing scientific consensus
that by the year 2050 global temperatures will have risen
significantly (Houghton ef al., 1990), with many studies
predicting warming of a magnitude not observed during
human history. Such climatic change could lead to large
impacts on individual organisms, communities, natural
ecosystems and global biochemical cycles and have
potentially grave impacts on biodiversity.
Large-scale patterns in the physiognomy and potential
species occurrence in different vegetation types are
primarily determined by climate. Climatic parameters, such
as temperature and precipitation, determine the major
boundaries between latitudinal zones (e.g. boreal, temperate
and tropical), and vegetation types (e.g. deserts, steppes and
forests). Temperature and precipitation, and their annual
variation, control the potential appearance of vegetation,
such as the distribution of deciduous or evergreen tree-
species, or short and long grass prairie-species. The
combined effects of climate, soil characteristics, vegetation
history, large-scale disturbances and anthropogenic
influences determine the actual vegetation both regionally
and locally.
This close correlation between climate and the physiognomy
of vegetation has for some time been recognised by
environmental scientists and has led to the use of vegetation
to create climate maps and vice versa (e.g. Koppen, 1936;
Holdridge, 1967). The Holdridge Life Zone Classification
(Holdridge, 1967; Plate 4) is often used for studies of the
impact of climate change. The ecoclimatic zones of this
model provide reasonable agreement with potential natural
vegetation patterns at a global scale. The Life Zone
Classification is based on the following annual climatic
variables: biotemperature (mean positive temperatures),
total annual precipitation and evaporation (defined as a
function of biotemperature). The Life Zones are delimited
by hexagons derived from a triangular graph of these three
variables. Maps of the Life Zone Classification can be
created for current climatic conditions (Plate 5) and for
potential conditions determined by climate change. To
obtain the best possible agreement with existing vegetation
patterns, the Life Zones have been aggregated into biomes
(large-scale vegetation assemblages).
MODELLING GLOBAL CLIMATE CHANGE
Global climate models can simulate the dynamics of the
atmosphere under different conditions. Such models can be
used, for example, to determine the potential climatic
change equivalent to a doubling of atmospheric carbon
dioxide. Detailed descriptions of these models and their
results can be found in Houghton et al. (1990). The results
used here are from the models of the Geophysical Fluid
Dynamics Laboratory (GFDL), Goddard Institute for Space
Studies (GISS), Oregon State University (OSU) and the
United Kingdom Meteorological Office (UKMO).
254
Although there are differences in the magnitude of change,
all models show similar patterns for a changed global
climate, particularly with respect to increased temperatures.
Greatest temperature increases occur during the winter
season in polar regions, and could exceed 15°C. The
pattern is less pronounced during the summer season, when
the overall temperature increase is less. The different
simulations agree less well in terms of precipitation
patterns. In general, the models predict a global increase in
precipitation, but there are large differences in the predicted
seasonal and regional patterns. Besides, many regions that
experience an increased precipitation could exhibit no
change or even a negative change in moisture availability
because of alterations in the balance between temperature,
precipitation and evapotranspiration. The models generally
predict a relatively modest rise in sea-level, unlikely to
exceed one metre over the present century. This rise would
be largely a result of thermal expansion of the oceans and
melting of minor ice-bodies rather than any major change
to the polar ice-caps.
EFFECTS OF CLIMATE CHANGE ON VEGETATION
ZONES AND BIODIVERSITY
Aggregated Life Zone Classifications have been generated
using the simulated climate-change scenarios (Plate 6: the
GISS model). Comparison of this map and Plate 5, for
current conditions, clearly displays the potential changes in
global vegetation patterns. Large changes in the current
extent and location of global vegetation zones are projected
and the different scenarios all show a similar pattern of
change (Plates 7 and 8). The changes are not consistent
across the globe but depend on the non-linear change in
both temperature and precipitation. Shifts of biomes are
most apparent in the mid and high latitude regions, with
only slight changes in the tropics. The boreal and polar
biomes show the largest polewards shift, with a decrease in
the extent of tundra and forested tundras. These biomes
currently form a continuous circumpolar band but under a
warmer climate only scattered patches remain. In
comparison, the current extent of tropical forests is rather
stable, with the total potential area of forest increasing. Any
actual increase in the tropical forests, however, will be
significantly constrained by human land-use and therefore
cannot be expected to evolve to the potential mapped extent.
The maps presented in this assessment give a general
indication of expected changes in the distribution of
ecoclimatic zones on a global scale. The specific impacts of
these changes on global biodiversity are difficult to assess
definitively at present, because little is known of the
physiological tolerance and potential migration capability of
numerous species. However, a preliminary illustration of
the potential threats these climatic changes could inflict on
biodiversity protection can be given by assessing the
impacts of ecoclimatic changes on a global distribution of
a selection of large (>1,000ha) nature reserves.
Biodiversity protection in a changing environment will be
influenced by both the magnitude and speed of
environmental change and also the ability of species to
respond to this change. In these terms, the effects of climate
change must be viewed in the context of natural ecosystem
fragmentation which may inhibit the migration of species to
more suitable habitats under future climates. When the
correlative ecoclimatic mapping presented above is overlaid
onto a global distribution of existing nature reserves,
numerous sites are shown to experience shifts in ecoclimatic
types. The climatic conditions normally associated with the
vegetation structure of many of these sites would be
expected to shift beyond the stationary boundaries of the
established reserves. Table 19.1 gives the percentage of a
selection of 2,618 nature reserves that are in areas affected
by large shifts in ecoclimatic zones under different climate
scenarios, marked as either ‘stable’ or ‘endangered’. It
should be noted that the inherent robustness of individual
reserves, for example those with a wide altitude range, has
not been considered in these data. The table also indicates
a CHANGE scenario which includes those reserves where
all climate change scenarios agree that there will be a
change from one life zone to another (cf. Plate 7). The
SIMILARITY scenario further demands that all four models
predict similar new life zones for the reserves concerned
(cf. Plate 8).
Table 19.1
Predicted life zone changes
in selected reserves
CLIMATE- STABLE ENDANGERED PERCENT
SCENARIO ENDANGERED
GFDL 1295 1323 50.5
GISS 1442 1176 44.9
OSU 1522 1096 41.9
UKMO 1097 1521 58.1
CHANGE 1754 864 33.0
SIMILARITY 2173 445 17.0
This translocation of ecoclimatic ranges could act to
fragment habitats further as species individually respond to
255
Global Climate Change
climatic changes. Local extinctions could occur through
either direct physiological responses to climatic conditions
or through changes in interspecific competition owing to
alterations in the composition and population of different
species groups within reserves (Peters and Darling, 1985;
Hunter et al., 1988). Changes in the future composition of
protected habitats may also have significant impacts beyond
the regional scale. Migratory species which exploit different
biomes seasonally or at different stages of their life histories
could be significantly affected by the climatic disruption of
reserve sites which link migration corridors or flyways.
This ‘biodiversity’ assessment depicts shifts in climatic
zones and links them with large nature reserves. The shifts
were interpreted as having an ecologically significant impact
on many reserve sites (Table 19.1). This assessment should,
however, not be used as a direct evaluation of the potential
loss or gain in biodiversity. However, the percentage of
impacted reserves indicates that the current system may not
provide the environmental requirements of many species
and ecosystems in the near future and will thus be less
capable of safeguarding biodiversity.
References
Holdridge, L.R. 1967. Life Zone Ecology. Tropical Science Center,
San José. 206pp.
Houghton, J.T., Jenkins, G.J. and Ephraums, J.J. (Eds) 1990. Climate
change: the IPCC scientific assessment. Cambridge University
Press, Cambridge. 365pp.
Hunter, M.L., Jacobson, G.L., Jr. and Webb, T. TI 1988.
Paleoecology and the coarse filter approach to maintaining
biodiversity. Conservation Biology 2:375-385.
KGppen, W., 1936. Das geographische System der Klimate. In:
K6ppen, W. and Geiger, R. (Eds) Handbuch der Klimatologie.
Berlin. 46pp.
Peters, R.L. Il and Darling, J.D.S. 1985. The greenhouse effect and
nature reserves. BioScience 35:707-717.
Contributed by Rik Leemans, Global Change Department,
National Institute of Public Health and Environmental
Protection, the Netherlands, and P.N. Halpin, Department
of Environmental Sciences, University of Virginia,
Charlottesville, Virginia, USA.
1. Biological Diversity
20. TROPICAL MOIST FORESTS
WHAT ARE TROPICAL MOIST FORESTS?
The terms ‘rain forest’ and ‘tropical moist forest’ are often
used as synonyms; although neither has a standard
definition, the latter is more inclusive than the former.
Schimper first used the term rain forest in 1903 (Schimper,
1903) and defined it as a forest that is "evergreen,
hygrophilous in character, at least 30m high, rich in thick-
stemmed lianas and in woody as well as herbaceous
epiphytes". Sixty or so years later, Baur (1964) extended
this definition somewhat to "a closed community of
essentially but not exclusively broadleaved evergreen
hygrophilous trees, usually with two or more layers of trees
and shrubs with dependent synusiae of life forms such as
vines and epiphytes. It includes the characteristic vegetation
of the humid tropics, even when this has a somewhat
seasonal climatic regime, as well as those of moist elevated
areas of the tropics".
In the following discussion of existing areas of tropical
forest and rates of change in cover, many of the data have
come from FAO and refer to ‘closed broadleaved forests’
which are again defined differently, and in particular
include dry, deciduous forests. The most comprehensive
atlas of tropical forests, two volumes of which have been
used in compiling Tables 20.9 and 20.10 (Collins er al.,
1991; Sayer et al., 1992), includes mangroves and montane
forests in the estimates of tropical moist forest, as well as
monsoon forests in Asia; they do not include riverine
forests or dry deciduous forests. The third volume in this
series, on Latin America, is still in preparation, hence the
lack of comparable data for that region in the cited tables.
Maps from this series, the most consistent and current
available, are reproduced in simplified form in Figs 20.10-
12. Many of the difficulties encountered in compiling
standard statistics of forest area arise from the use of
different or inconsistent definitions of vegetation type.
The significance of tropical forests
The forests are home to millions of people, providing them
with shelter, food, clothing, fuel, medicines, building
materials and a variety of other resources. They are also the
origin of many of these same resources for countless people
who do not actually live in the forest.
The commodity that is generally considered to be of the
greatest economic value is timber. However, the
commercial value of other products such as fruits, nuts,
rattans, medicinal plants and rubber - which can be cash
crops or for local use - are frequently not taken into account
(Peters et al., 1989). Southeast Asia, in particular, has a
long history of successful export of non-timber forest
products such as rattans, resins and gums (Reitbergen,
1992). Latin America’s main non-timber forest exports have
been rubber and brazil nuts. In general, non-timber
commodities appear to be less significant among forest
products of Africa.
Numerous species important to pharmaceutical companies
are derived from the rain forests at present and it is
predicted that many more will be found if time and money
is invested in the search for them. Local people use the
forest products to a considerable extent in treating their own
ailments and these can form a starting point for
investigation by others.
Apart from producing many resources of subsistence and
commercial importance, the forests play a key role in
regulating water flow, conditioning local climate and
protecting against soil erosion. Although the role they play
in influencing local rainfall is not well-understood, it is
clear that this can be significant. In several places where
forests have been destroyed there has been a reduction in
rainfall. For instance in Banjul, the capital city of The
Gambia, in 1965 when there was still good forest cover,
annual precipitation was 1,240mm. Between 1982 and
1988, when the forest had all but disappeared, the mean
level was almost halved to 650mm (Jones, 1992). There is
reasonable evidence that reduction in rainfall can be a
consequence of forest clearance. Preservation of the tropical
rain forests is also vital for conserving biodiversity.
Although they cover only 6-7% of the earth’s surface, these
forests probably contain more than 50%, and possibly as
much as 90%, of all species of plants and animals.
Factors leading to tropical forest degradation
The timber trade is widely considered to be responsible for
much of the destruction of the rain forests, partly directly,
but mainly indirectly, by opening up formerly pristine areas
to invasion by shifting cultivators. Mining and oil
companies have the same effect, leaving roads into the
forest and attracting settlers to an area. All three industries
cause direct damage as well. Logging can severely degrade
an area if not done selectively and with care, as is likely to
occur when companies have no stake in the long-term
sustainability of supplies. Pollution from mining or oil
drilling gives rise to further problems. In some cases, vast
quantities of fuel are needed to power a mining programme
or other industry and this can be responsible for further
devastation. For instance, iron-ore smelters in the Brazilian
Grand Carajas Programme will consume 2,300km* of forest
as charcoal each year.
Though not threatening the forest directly, invasion by
commercial companies also displaces indigenous peoples,
and this is a major cause of concern in South America and
Southeast Asia. The building of dams has also resulted in
large areas of forest being lost through flooding and, more
importantly, this has often caused major ecological
problems in nearby areas as well as encouraging road
development and settlement.
It is considered by many that the most important agent of
tropical forest destruction is the shifting cultivator. Poverty,
population growth and unequal land ownership are the
fundamental causes of this form of land conversion. In
many cases, governments encourage peasants from high
population areas to move into less developed, usually
forested, areas (see case studies on forest destruction in
Rondonia and transmigration in Indonesia). Some forest
destruction, particularly in Brazil and Central America, has
occurred as a result of the tax incentives offered to those
who cleared forest for cattle ranches.
THE GLOBAL AREA OF TROPICAL
FOREST
MOIST
There are almost as many estimates of the present extent of
tropical forests and rates of deforestation as there are
reports about the subject. The problems associated with
trying to obtain accurate figures for how much forest cover
exists today are greatly multiplied when considering the
cover that existed 20 or 50 years ago.
There are two major problems to be overcome before any
attempt can be made at calculating either forest area (past
or present) or rate of deforestation, and these are essentially
problems of definition. The term ‘forest’ has to be defined
and applied consistently throughout the study; this is
frequently not done and such variation has given rise to
great differences in the estimates of forest cover in some
countries. The second problem is that some authors have
taken ‘deforestation’ to mean the complete destruction of a
forest while others have included areas that have been
degraded (by logging in particular), and have thereby
estimated much higher rates of deforestation. Increased
precision over the definition and application of these terms
is desirable, and clarification as to which are being included
in the estimate of deforestation. It is generally very difficult
to quantify degradation of an area and yet this type of
disturbance can have a significant and protracted effect on
the species composition (both flora and fauna), biomass and
structure of a forest.
One of the first comprehensive estimates of how much
tropical forest existed was made by Sommer (1976) who
noted that "a global appraisal of tropical moist forests
undertaken at this time can only base its research on the
material available - a mass of incomplete data and a number
of assumptions. It will yield rather rough results."
Sommer defined what categories of tropical forest he was
including in his estimates and which countries he was
covering within his regional reviews. Estimates of climax
areas of tropical moist forests were taken from vegetation
maps and these sources and their problems are all reported.
Tropical Moist Forests
Figures for present areas of moist tropical forest were
calculated from information at FAO headquarters. The main
sources of data were land-use and vegetation maps, project
reports and country statistics, occasionally supplemented by
oral reports. For each country under consideration some
detail on the problems encountered are presented, and
Sommer makes it quite clear that he is not providing
definitive figures from reliable data. He concluded that at
the time of his research there were 9,350,000km* of
tropical moist forest in the world (Table 20.1).
Sommer gives a list of 13 countries for which he was able
to find figures in "various reports" for the area of forest
lost per year. The character of the clearing was in most
cases not reported. From this figure, of 21,600km?, he
extrapolated to all countries with tropical moist forest and
obtained an estimated deforestation rate of 110,000km? per
year.
Sommer concludes his report by noting that "an accurate
appraisal of the climax and actual areas of the moist tropical
forest at the global level, based on the information available
at FAO headquarters, is not yet possible".
In 1980, Myers wrote a report for the National Academy of
Sciences on Conversion of Tropical Moist Forests. He too
noted that "sound information, especially authoritative
statistical information, is not easily obtained". The data
sources used by Myers were mostly published reports,
generally limited to those produced in the 1970s, combined
with correspondence, discussions and visits to three
Southeast Asian and three South American countries.
He noted that there is no standard and objective
classification of ‘tropical moist forest’ (TMF) but, after
consulting numerous sources, he uses the definition
"evergreen or partly evergreen forests, in areas receiving
not less than 100mm of precipitation, in any month for two
out of three years, with mean annual temperatures of
24+°C and essentially frost-free; in these forests some
trees may be deciduous; the forests usually occur at
Table 20.1 _ Areas of tropical moist forest estimated by Sommer (1976)
REGION PRESENT
MOIST
FOREST AREA
(thousand km?)
Africa
East Africa 70
Central Africa 1,490
West Africa 190
Total 1,750
Tropical America
Latin America 4,720
Central America and Caribbean 340
Total 5,060
Asia
Pacific 360
Southeast Asia 1,870
South Asia 310
Total 2,540
TOTAL HUMID TROPICS 9,350
PER CENT PER CENT
OF WORLD’S OF TOTAL
MOIST LAND AREA
FOREST
0.7 10.6
15.0 65.9
2.0 19/1
18.7 36.2
50.5 53.5
3.6 31.9
54.1 51.2
3.8 12.8
20.0 67.4
3.3 24.4
27.2 37.2
100.0 42.8
Source: Sommer, A. 1976. Attempt at an assessment of the world’s tropical moist forests. Unasylva 28(112-113):5-24.
257
1. Biological Diversity
altitudes below 1,300m (though often in Amazonia up to
1,800m and generally in Southeast Asia up to only 750m);
and in mature examples of these forests, there are several
more or less distinctive strata". However, as several other
authors have noted (e.g. Lugo and Brown, 1982; Holdgate,
1982; Sayer and Whitmore, 1991), he does not
subsequently use this definition when giving forest areas for
some of the countries discussed.
In his chapter on the role of forest farmers, Myers (1980,
p.25) states that "it is not unrealistic to suppose that forest
farmers are converting at least 100,000km? of primary
forest to permanent cultivation each year". He then
continues "when considered in conjunction with other
factors - timber harvesting, planned agriculture, cattle
raising, etc. - it becomes possible to credit that something
approaching 200,000km? of TMF, and possibly even more,
are being converted each year". He noted earlier that
conversion can range from marginal modification to
fundamental transformation, and it is not clear how drastic
a conversion the "other factors" are causing.
Myers’ report continues with regional reviews for Southern
and Southeast Asia including Melanesia (14 countries and
Melanesia); Tropical Latin America (18 countries and the
Caribbean); and Tropical Africa (13 countries). The degree
of detail given for each country is very variable and in most
instances the causes of deforestation are concentrated on,
rather than the rate. No comprehensive summary of
deforestation rates or forest area in each country is
presented and, in many cases, this information cannot be
extracted from the text. However, the report does identify
areas particularly at risk from deforestation and its intention
was probably never to estimate forest areas but to document
the different forms and degrees of conversion of tropical
moist forest that were taking place.
The most comprehensive reports on forest cover and rates
of deforestation are those produced by FAO/UNEP in 1981
and updated in 1988. The 1981 study was carried out on a
national basis, and this information was then used to
compile a regional synthesis for each of the three tropical
areas (Tropical Africa, Tropical Asia and Tropical
America) from which a global assessment of forest cover
and rate of deforestation was derived. An FAO Forestry
paper (Lanly, 1982) was subsequently produced which
collated all the findings of the Tropical Forest Resources
Assessment Project in an overall synthesis for the tropical
world.
Forest extent was estimated for each of the 76 countries (23
in tropical America, 37 in tropical Africa and 16 in Asia)
covered by the project using the best available sources of
information. The whole project occupied a total of almost
seven man years. Data collection involved visits to research
institutes in Europe, particularly those involved in the study
and mapping of vegetation, visits to national forestry, land-
use and survey departments and to regional FAO offices,
considerable correspondence with the forestry services, use
of satellite imagery for 18 countries and interpretation of
satellite imagery for the 13 countries where no other
information was available. In three cases the assessments
were carried out by the countries themselves. Rates of
forest removal were taken either from the degree of shifting
258
cultivation or land-use statistics. These rates were used to
extrapolate forest cover to a common baseline of December
1980. Estimates for deforestation rates for 1981-1985 were
also made but these projections can be only broadly
indicative of trends and future conditions and, as the report
states, they have to be viewed with caution.
As a result of the differences in information quality,
estimates of forest cover and deforestation rates are judged
to be very reliable for only 15 of the 76 countries surveyed.
In terms of total area of closed tropical forests, however,
this represents 40% of forest extent, largely because Brazil
is in the very reliable group. A further 38 countries
(covering another 40% of closed forest) have very good
baseline data on forest cover while in the remaining 23
countries, both baseline data and deforestation rates are
considered to be of medium to poor quality.
The classification of vegetation types used by FAO/UNEP
has placed particular emphasis on forest management. In the
update of these reports (FAO, 1988), which includes a
further 53 countries, a clear distinction has been made
between the open and closed forests in the tabulated data
provided. The closed broadleaved forests, which are the
ones that are usually equated with the term tropical moist
forest, are defined as "generally, but not always multi-
storeyed, and may be evergreen, semi-deciduous or
deciduous and occur in wet, moist or dry zones. They cover
with their various storeys and undergrowth, a high
proportion of the ground; and do not contain a continuous
dense grass layer" (FAO, 1988).
The open forests, in contrast, “correspond to mixed forest-
grassland formations, with a continuous dense grass layer
in which the crowns of the trees cover more than 10% of
the ground. They thus include e.g. the various forms of
cerrado and chaco in America; and tree and wooded
savannas and woodlands in Africa" (FAO, 1988).
In FAO’s 1988 summary report, information on areas of
bamboo and coniferous formations is also supplied, and
each forest type is broken down into operable forests
(which may be unmanaged - either virgin or logged - or
managed) or inoperable forests (for either physical or legal
reasons). Information on deforestation is similarly presented
within these classes of forest type and management status.
The 1981 study (Lanly, 1982) concluded that in 1980 there
were approximately 12 million km? of closed forest, of
which 97% was closed broadleaved, and 7,350,000 km? of
open tree formations remaining in the tropical world (Table
20.2).
The 1988 updated report indicated that about 75,000 km? of
closed forest (Table 20.3) and of open formations (Table
20.4) were being cleared each year between 1981 and 1985
to allow the land to be used for other purposes. This is an
average reduction rate of 0.62% for the closed forest and
0.52% for the open formations. In addition to this
deforestation, considerable degradation was occurring,
particularly in the open forest, caused by overgrazing,
fuelwood gathering and repeated burning. This, however,
is more gradual and difficult to quantify.
Tropical Moist Forests
Table 20.2. FAO/UNEP (1981) estimates of the area of tropical forest remaining in
1980
CLOSED FOREST' OPEN FOREST
(km?) (km?)
Tropical America 6,786,550 2,169,970
Tropical Africa 2,166,340 4,864,450
Tropical Asia 3,055,100 309,480
WORLD 12,007,990 7,343,900
Source: Lanly, J.-P. 1982. Tropical Forest Resources. FAO Forestry Paper 30, FAO, Rome, Italy.
Note: ' Includes closed broadleaved, coniferous and bamboo forest.
Table 20.3. FAO(1988) estimates of annual areas deforested and deforestation rates
in closed forests’
TOTAL AREAS DEFORESTED (km?) DEFORESTATION RATES (per cent)
1976-1980 1981-1985 1976-1980 1981-1985
Americas 41,190 43,390 0.60 0.63
Africa 13,330 13,310 0.61 0.61
Asia 18,150 18,260 0.59 0.60
TOTAL 72,670 74,960 0.60 0.62
Source: FAO 1988. An Interim Report on the State of the Forest Resources in the Developing Countries. FAO, Rome, Italy.
Note: ' Includes closed broadleaved, bamboo and coniferous forests.
Table 20.4 FAO (1988) estimates of annual areas deforested and deforestation rates
in open forests
ANNUAL AREA DEFORESTED DEFORESTATION
(km?) RATES (per cent)
1981-1985 1981-1985
Americas 12,720 0.59
Africa 23,450 0.48
Asia 1,900 0.61
TOTAL 38,070 0.52
Source: FAO 1988. An Interim Report on the State of the Forest Resources in the Developing Countries. FAO, Rome, Italy.
Another source of figures on extent of forests in the tropics should be published in late 1992. There have, however,
is the land-use tables in the FAO Production Yearbooks. already been two interim reports (FAO, 1990, 1991) and an
These give data over time for each country on the extent of evaluation of the first of these (Lanly er al., 1991).
arable land, permanent crops, permanent pasture, forest and
woodland, and other land. The area of forest and woodland In both interim reports, forests are defined as "ecological
"refers to land under natural or planted stands of trees, systems with a minimum of 10% crown cover of trees
whether productive or not, and includes land from which and/or bamboos, generally associated with wild flora and
forests have been cleared but that will be reforested in the fauna and natural soil conditions and not subject to
foreseeable future." This is not, therefore, a particularly agricultural practices". Deforestation refers to "change of
meaningful figure, and especially not when data on moist land use or depletion of crown cover to less than 10%".
forests alone are required. However, the increase in the The first interim report gives provisional estimates of forest
area under crops over time can, in some countries, be an cover and deforestation for 62 countries, mostly in the
indication of how much forest is being converted to moist tropical zone (Table 20.5). It is stressed that parallel
agricultural land so some authors (e.g. Mather, 1990) have developments cannot be assumed to have taken place in the
been able to use these data in their reports on deforestation. forests of the dry and mountainous zones.
FAO is in the process of updating its information on each
country and region to produce a second detailed report on By the time the second interim report was published, data
forest resources with 1990 as the reference year. This on a further 25 countries had been included and existing
259
1. Biological Diversity
Table 20.5 Preliminary FAO (1990) estimates of 1990 forest area and deforestation
for 62 countries in the tropics
CONTINENT NO. OF TOTAL FOREST FOREST ANNUAL DEFOREST.
COUNTRIES LAND AREA AREA DEFOREST. RATE
STUDIED AREA 1980 1990 1981-90 1981-90
(km?) (km?) (km?) (km?) (%)
Africa 15 6,098,000 2,897,000 2,418,000 48,000 1.7
Latin America 32 12,636,000 8,259,000 7,530,000 73,000 0.9
Asia 15 8,911,000 3,345,000 2,875,000 47,000 1.4
TOTAL 62 27,645,000 14,501,000 12,823,000 168,000 ibys
Source: FAO 1990. Interim Report on Forest Resources Assessment 1990 Project. Committee on Forestry Tenth Session. FAO, Rome, Italy.
Table 20.6 Preliminary FAO (1991) estimates of forest area and deforestation for 87
countries in the tropics
CONTINENT NO. OF TOTAL FOREST FOREST ANNUAL DEFOREST.
COUNTRIES LAND AREA AREA DEFOREST. RATE
STUDIED AREA 1980 1990 1981-30 1981-90
(km?) (km?) (km?) (km?) (%)
Latin America 32 16,756,000 9,229,000 8,399,000 84,000 0.9
Central America & Mexico 7 2,453,000 770,000 635,000 14,000 1.8
Caribbean Sub-region 18 695,000 488,000 471,000 2,000 0.4
Tropical South America 7 13,608,000 7,971,000 7,293,000 68,000 0.8
Asia 15 8,966,000 3,108,000 2,748,000 35,000 1.2
South Asia 6 4,456,000 706,000 662,000 4,000 0.6
Continental SE Asia 5 1,929,000 832,000 697,000 13,000 1.6
Insular SE Asia 4 2,581,000 1,570,000 1,389,000 18,000 1.2
Africa 40 22,433,000 6,504,000 6,001,000 51,000 0.8
West Sahelian Africa 8 5,280,000 419,000 380,000 4,000 0.9
East Sahelian Africa 6 4,896,000 923,000 853,000 7,000 0.8
West Africa 8 2,032,000 552,000 434,000 12,000 2.1
Central Africa 7 4,064,000 2,301,000 2,154,000 15,000 0.6
Tropical Southern Africa 10 5,579,000 2,177,000 2,063,000 11,000 0.5
Insular Africa 1 582,000 132,000 117,000 2,000 1.2
TOTAL 87 48,155,000 18,841,000 17,148,000 170,000 0.9
Source: FAO 1991. Second Interim Report on the State of Tropical Forests by Forest Resources Assessment 1990 Project. Tenth World Forestry
Congress, September 1991, Paris, France.
Table 20.7 A comparison of forest area and deforestation rate as estimated in FAO’s
tropical forests resource assessment projects for 1980 and 1990
PROJECT FOREST AREA ANNUAL AREA DEFORESTATION
DEFORESTED RATE
(km?) (km?) (%)
Reference year 1980 19,350,000 113,000 (for 1981-85) 0.6
Reference year 1990 18,820,000 169,000 (for 1981-90) 0.9
information had been updated. Although deforestation rates
for Latin America and Asia remained similar to those
presented in the 1990 report, the rate of change given in
this report is considerably lower for Africa (Table 20.6),
presumably because of the inclusion of vast areas of open
forest. It must be noted that in both these interim reports,
no distinction has been made between open and closed
260
forests whereas, in the discussions above, figures from the
FAO/UNEP (1981) report are for closed forests only.
When a comparison is made between the FAO assessments
for 1990 and 1980 using the data from the 76 countries
common to the two reports, it is apparent that the annual
rate of deforestation has risen considerably (Table 20.7).
Tropical Moist Forests
Deforestation estimates for closed tropical forests’, for selected countries
RECENT NON-FAO ESTIMATES
ANNUAL AREA ANNUAL RATE PERIOD OF
LOST OF Loss RECENT
(km?) (%) ESTIMATES
80,000 ~ 2.2 1987
35,000 e 1987
1,000 ; 0.6 1976-86
1,000 2 c. 1987
1,240 : 7.6 1977-83
6,000 d 1960-84°
15,000 i 41 1975-82
1,500 : c. 1987
9,000 0.8 1979-84
10,000 +" c. 1985
677000) 2.1 1975-81
3,000 1“ 1988
1,430 1“ 1.5 1981-88
1.0 ” 1988
a7) 2.5 1978-85
3700 c. 1985
1730 ee 2.0 1976-81
Table 20.8
FAO ESTIMATES 1981-85'>
COUNTRY ANNUAL AREA ANNUAL RATE
LOST OF LOSS
(km?) (%)
Brazil 14,800 0.4
Cameroon? 800 0.4
Costa Rica 650 4.0
Colombia 8,200 0.4
India® 1,470 0.3
Gabon 150 0.1
Indonesia 6,000 0.5
Myanmar 1,050 0.3
Peru 2,700 0.4
Philippines 920 1.0
Thailand! 3,790 2.4
Malaysia 2,550 ee
Viet Nam 650 0.7
Sources:
' FAO 1988. ? Setzer, A.W. et al. 1988. * Fearnside, P.M. 1990. * Joint Interagency Planning and Review Mission (JIM) 1988.
* FAO/UNDP 1988. ° Sader and Joyce 1988. 7 Plan de Accién Forestal de Colombia, perfil de proyectos (undated). * Vohra, B.B. 1987. ° IUCN
1989. ' The World Bank 1988. '! USAID 1987. ' Kyaw, U.S. 1987. '° Saavedra, C. and Suarez de Freitas, G. 1989. '* Philippines Forest
Management Bureau 1988. '° Forest Management Bureau 1988. ' Royal Forestry Department of Thailand 1986. '’ Thang, H.C. 1987. '* Vo
Quy 1988.
Notes: * Closed forests are forests in which trees cover a high proportion of the ground and in which grass does not form a continuous layer on
the forest floor. Open forests are forests in which trees are interspersed with grazing lands. » Unless otherwise noted, annual deforestation rate is
calculated from FAO 1981 estimates. © For Legal Amazon only. Brazil also has a small amount of closed coastal forest remaining. * Annual
deforestation rate is calculated from data found in sources 4 and 8, above. ° This long period over which deforestation rate has been estimated may
be the cause of the comparatively low figure given. ‘ Represents total forests, open and closed.
FAO (1990) and Lanly et al. (1991) give three possible
reasons for this increase in the rate of deforestation between
1980 and 1990. These are:
©@ an actual increase of rate of deforestation
® an underestimation of the rate of deforestation in the
1980 assessment
©@ an overestimation of the rate of deforestation in the 1990
assessment.
At this stage, FAO has not been able to assess the relative
contribution of these various components. It is hoped,
however, that the final results, based on uniform remote
sensing observations made specifically for the 1990 project,
will provide a more accurate estimate of forest cover which
can then be used in calculations of deforestation rates.
Reports other than those by FAO have also indicated that
there has indeed been an increase in the rate of
deforestation over the past decade. WRI (1990) reported a
considerably higher rate of deforestation for the eight
countries for which they obtained figures compared with the
estimates of FAO (1988). Their data, presented in Table
20.8, are for closed forests (except in Thailand) and, if
accurate, suggest that around 204,000km? of this forest type
are being lost annually. Sayer and Whitmore (1991) give a
table comparing annual rates of deforestation as estimated
by FAO (1988) for 1981-1985 and those given by Myers
(1989). These, together with other rates of deforestation
from more recent sources, have been incorporated in Table
20.8. In all countries except Colombia, FAO gives lower
estimates than the other sources report. [IUCN and WCMC
are in the process of compiling a series of atlases showing
the extent of forest present today. The first volume, on Asia
and the Pacific has been published, the second, on Africa
is in press, while the third, on Latin America and the
Caribbean, is in preparation. Data from the first two
volumes have been used in compiling Tables 20.9 and
20.10 and simplified maps derived from the series are
1. Biological Diversity
Table 20.9 Tropical moist forests: original versus remaining extent
APPROXIMATE REMAINING EXTENT OF MOIST FORESTS (KM) % MOIST FOREST REMAINING
ORIGINAL EXTENT OF From aties Publication FAO (1988) data From map From FAO
CLOSED CANOPY maps; rain & date of maps for 1980, closed data (1988)
TROPICAL MOIST monsoon broadleaved plus data
FORESTS (KM?) forests coniferous forests
ASIA
Bangladesh 130,000 9,730 1981-6 9,270 7.0 7.0
Brunei 5,000 4,692 1988 3,230 94.0 65.0
Cambodia 160,000 113,250 1971 71,680 71.0 45.0
China/Taiwan 340,000 25,860 1979 125,860 8.0 =
India 910,000 228,330 1986 504,010 25.0 55.0
Indonesia 1,700,000 1,179,140 1985-9 1,138,950 69.0 67.0
Laos 225,000 124,600 1987 78,100 55.0 35.0
Malaysia 320,000 200,450 - 209,960 63.0 66.0
Peninsular (130,000) (69,780) 1986 = 54.0 =
Sabah (70,000) orang 1984 = 51.0 -
Sarawak qi 70000) (94,670) 1979 = 79.0 =
Myanmar 600,000 311,850 1987 313,090 52.0 52.0
Philippines 295,000 66,020 1988 95,100 22.0 32.0
Singapore 500 20 (1980s) = 4.0 -
Sri Lanka 26,000 12,260 1988 16,590 47.0 64.0
Thailand 250,000 106,900 1985 83,350 43.0 33.0
Viet Nam 280,000 56,680 1987 75,700 20.0 27.0
OCEANIA
Australia 11,000 10,516 1988 10,516 96.0 -
Fiji 18,000 6,970 (1980s) 8,110 39.0 45.0
Papua New Guinea 450,000 366,750 1975 342,300 82.0 76.0
Solomon Islands 27,500 25,590 (1980s) 24,230 90. 90.0
AFRICA
Angola 218,200 = = 29,000 = 13.3
Benin 16,800 424 1989-90 and 79 470 2.5 2.8
Burundi 10,600 413 1984 150 3.9 1.4
Cameroon 376,900 155,330 1985 179,200 41.2 475
Central African Rep 324,500 52,236 1985 35,900 16.1 11.1
Comoros 2,230 - - 160 = 7A
Congo 342,000 = - 213,400 = 62.4
Cote d'Ivoire 229,400 27,464 1989-90 44,580 12.0 19.4
Djibouti 300 = ce a 4 38
Equatorial Guinea 26,000 17,004 1960 12,950 65.4 49.8
Ethiopia 249,300 = = 27,500 = 11.0
Gabon 258,000 227,500 = 205,000 88.2 79.5
Gambia 4,100 497 1985 650 124 15.6
Ghana 145,000 15,842 1989-90 17,180 10.9 11.8
Guinea 185,800 7,655 1989 20,500 41 11.0
Guinea-Bissau 36,100 = = 6,660 = 18.4
Kenya 81,200 = = 6,900 = 8.5
Liberia 96,000 41,238 1989-90 20,000 43.0 20.8
Madagascar 275,086 41,715 1985 103,000 15.2 37.4
Malawi 10,700 320 = 1860 3.0 17.4
Mauritius 1,850 - - 30 = 1.6
Mozambique 246,900 - - 9,350 - 3.9
Nigeria 421,000 38,620 1989-90 59,500 9.2 141
Reunion 2,500 - - 820 = 32.8
Rwanda 9,400 __ 1,554 (nd) 1,010 16.5 10.7
Sao Tomé and Principe 960 209 1985 560 31.1 58.3
Senegal 27,700 2,045 1985 2,200 74 7.9
Seychelles 270 = = 30 = 11.41
Sierra Leone 71,700 5,064 1989-90 7,400 7A 10.3
Somalia 21,200 = = 14,800 = 69.8
Sudan 27,000 - - 6,400 = 23.7
Tanzania 176,200 = = 14,400 = 8.2
Togo 18,000 1,360 1989-90 3,040 76 16.9
Uganda 103,400 7,400 - 7,500 7.2 7.3
Zaire 1,784,000 1,190,737 1990 1,056,500 66.7 59.2
Zimbabwe 7,700 80 - 2,000 1.0 26.0
Notes: The data for Asian countries in column 1 are adapted from IUCN 1986. Review of the Protected Areas System in the Indo—Malayan Realm. 1UCN,
Gland. In the absence of comparable data for Australia and southern China, the totals in column 4 have been calculated using the map figures in
column 2. The FAO data for India are not directly comparable with the map data, as the former includes India’s extensive thorn forests. [The remaining area
figure for Cambodia in column 3 is now known to be 71,500km? according to FAO 1988. An Interim Report on the State of Forest Resources in the
Developing Countries. FAO, Rome.] Data in column 1 are mostly from MacKinnon, J. and MacKinnon, K. 1986. Review of the Protected Area
System in the Afrotropical Realm. (UCN, Gland; (except for Gabon and Liberia which were too high). Where this source gave no estimates for forest cover
of small islands it was assumed that they were once totally forested. The figures for Equatorial Guinea and Zaire in column 2 include 7,945km? and
86,547km? of degraded lowland rain forest respectively. The figure for Gabon in column 2 is from IUCN 1990. La Conservation des Ecosystémes Forestiers
d'Afrique Centrale. UICN, Gland. The figure in column 2 for Madagascar has been calculated by adding the figure from Green, G.M. and Sussman,
R.W. 1990. Deforestation history of the eastern rain forests of Madagascar. Science 248:212— 215; for eastern rain forest to that calculated for mangroves
from Map 26.1 in Africa plus an estimated 400km? for forest remaining in the Sambirano region. The figure for Malawi in column 2 is from
Dowsett—Lemaire, F. 1989. The flora and phytogeography of the evergreen forests of Malawi. I: afromontane and mid—altitude forests. Bulletin du Jardins
Botanique National de Belgique 59:3—131; and Dowsett—Lemaire, F. 1990. The flora and phytogeography of the evergreen forests of Malawi. II: lowland
forests. Bulletin du Jardins Botanique National de Belgique 60:9— 71. The figure in column 2 for Uganda is from Howard, P.C. 1991. Nature Conservation in
Uganda's Tropical Forest Reserves. IUCN, Gland. The figure for Zimbabwe in column 2 was supplied by T. Muller, in litt. The digital dataset in
column 3 for Zaire was completed in 1990 but is based on 1988 data. — no data available. (nd) = no date.
Tropical Moist Forests
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263
1. Biological Diversity
Notes for ASIA section of the Table 20.10 Tropical Moist Forests (Protected Area Coverage).
Figures given in column 2 are derived from maps in chapters 12-29 of Collins, N.M., Sayer, J.A. and Whitmore, T. 1991. The Conservation Atlas
of Tropical Forests: Asia and the Pacific. TUCN, Gland; the data varies in age.
The data in columns 3-6 for India refers only to the Western Ghats, north-east India and the Andaman and Nicobar Islands. There are no tropical
rainforests outside of these regions, but monsoon forest is extensive.
The totals given in columns 4,5 and 6 are for protected areas of greater than SOknr in extent which contain at least some tropical moist forest.
The totals in column 6 are derived from those protected areas which contain tropical forest mapped in chapters 12-29 of Collins, N.M., Sayer, J.A.
and Whitmore, T. 1991. The Conservation Atlas of Tropical Forests: Asia and the Pacific. TUCN, Gland.
The data for Australia refers only to tropical rain forest and other moist forest types are not included. Protected areas data refers only to national
parks.
The remaining area figure for Cambodia in column 3 is now known to be 71,500 kn? according to FAO 1988. An Interim Report on the State of
Forest Resources in the Developing Countries. FAO, Rome. 475pp.
Notes for AFRICA section of the Table 20.10 Tropical Moist Forests (Protected Area Coverage).
Data in column 2 is taken from White, F. 1983. The Vegetation of Africa: a descriptive memoir to accompany the UNESCO/AETFAT/ UNSO
vegetation map of Africa. Unesco, Paris; as in MacKinnon and MacKinnon, 1986, except for Gabon and Liberia where these authors indicate that
both countries were originally completely forested but the figures they give are country areas rather than land areas. "Original" cover includes
mosaics.
The figure in column 2 for Madagascar has been calculated by adding the figure from Green, G.M. and Sussman, R.W. 1990. Deforestation history
of the eastern rain forests of Madagascar. Science 248:212-215; for eastern rain forest to that calculated for mangroves from Map 26.1 in Collins,
N.M., Sayer, J.A. and Whitmore, T. 1991. The Conservation Atlas of Tropical Forests: Asia and the Pacific. TUCN, Gland; plus an estimated 400
km? for forest remaining in the Sambirano region.
The data given in column three are derived from maps in chapters 11-32 of Collins, Sayer and Whitmore (1991), unless stated otherwise.
The data in column 3 for Angola, Comoros, Congo, Djibouti, Ethiopia, Guinea-Bissau, Kenya, Mauritius, Mozambique, Reunion, Seychelles,
Somalia, Sudan, and Tanzania is from FAO 1988. An Interim Report on the State of Forest Resources in Developing Countries. FAO, Rome. 18pp.
The figure for Equatorial Guinea in column 3 includes 7,945 km? of degraded lowland rain forest.
The figure for Gabon in column 3 is from IUCN 1990. La Conservation des Ecosystémes Forestiers d'Afrique Centrale. UICN, Gland. 124pp.
The figure for Malawi in column 3 is from Dowsett-Lemaire, F. 1989. The flora and phytogeography of the evergreen forests of Malawi. I:
afromontane and mid-altitude forests. Bulletin du Jardins Botanique National de Belgique 59:3-131; and Dowsett-Lemaire, F. 1990. The flora and
phytogeography of the evergreen forests of Malawi. II: lowland forests. Bulletin du Jardins Botanique National de Belgique 60:9-71.
The figure for Uganda in column 3 is from Howard, P.C. 1991. Nature Conservation in Uganda's Tropical Forest Reserves. YUCN, Gland. 313pp.
The figure for Zimbabwe in column 3 supplied by T. Muller, in litt.
Data in columns 3-6 does not include forest reserves. Totals are for protected areas which contain at least some tropical moist forest as determined
on the maps in Collins, Sayer and Whitmore (1991), (except in Kenya, Ethiopia and Tanzania; see last note); it is not possible to take account of
fragmentation of forest within each protected area. In many cases the forest coverage will be over-optimistic.
* No data
Percentage of forest protected cannot be realistically calculated for Kenya, Ethiopia and Tanzania. This is because although there are protected areas
with forest within their boundaries, the forests are often fragmented and small in size and only cover a fraction of the size of the actual protected
area.
shown in Figs 20.10-12. ‘Forest’ in these atlases includes FACTORS INVOLVED IN CHANGES IN FOREST
mangroves and montane forests as well as lowland rain COVER
forests and swamp forests. In Asia, the monsoon forests are
also included in the statistics given, while in Africa dry Human occupation of forests dates back to 25,000-40,000
forests are excluded, as are riverine forests in both regions. years ago in Southeast Asia and the Pacific, 10,000 years
in the Amazon and perhaps 3,000 years in Africa (Poore
From Table 20.9 it can be seen that, in general, and Sayer, 1991). However, for most of man’s history, his
considerably less of the original forest extent remains in effect on tropical forests has been limited. Until
Africa than it does in Asia. For instance, nine of the 18 comparatively recently, populations densities were low and
Asian countries listed still have more than 50% of their there was little if any harvesting of trees for timber or
estimated original forest remaining while only four of the extensive clearing for agriculture. Changes were brought
36 African countries have this much of their original forest about when people moved from China into Southeast Asia,
left. There is also, overall, considerably more moist forest and from Europe to Africa and South America. Table
remaining in Asia than there is in Africa. Fig 20.13 20.11, compiled by Williams (1990), gives some indication
represents country area, forest area and annual loss in select of the areas of forest cleared in the tropics through time,
tropical countries (derived from FAO sources cited above). with a high and low estimate for each region. As he points
out though, there is much guesswork involved and the
Conclusions figures should not be taken as definitive. It is also not
entirely clear what formations are regarded as ‘forest’ in
It is apparent that the true extent of the remaining moist this calculation, but they almost certainly include dry forests
tropical forests is still unknown. However, in the past and possibly woodland where the canopy is not a closed
decade there have been marked advances in the capacity of one.
satellites to achieve detailed images of vegetation cover and
in the capacity of image interpreters to distinguish the From around 1600 the tropical forests were altered radically
different forest types (Myers, 1988). As a result, it should by the introduction of new crops and new methods of
soon be possible to obtain a more accurate assessment of exploitation (Williams, 1990). Forests were cleared to make
forest area. Similarly, rates of deforestation remain guesses way for cash crops such as rubber in Malaysia and
in many instances but whatever the figures, it is generally Indonesia, coffee in Brazil, tea in India and China, sugar in
agreed that they are high and are increasing. the Caribbean, tobacco and palm oil in Asia. In addition to
264
Table 20.11
REGION PRE-1650
Central America H 18
L 12
Latin America H 18
iL 12
Asia H 974
L 640
Africa H 226
E 96
Tropical Moist Forests
Estimated area of forest cleared in historical time (km? x 1,000)
1650-1749 1750-1849 1850-1978
30 40 200
30 40 200
100 170 637
100 170 637
216 596 1220
176 606 1220
80 -16 469
24 42 469
Source: Williams, M. 1990. Forests. In: Turner, B.L., Clark, W.C., Kates, R.W., Richards, J.F., Mathews, J.T. and Meyer, W.B. (Eds), The
Earth as Transformed by Human Action. Global and regional changes in the biosphere over the past 300 years. Cambridge University Press,
Cambridge.
Notes: H = high estimate, L = low estimate. Data for areas outside the tropics in the original table are omitted here.
crops, domestic animals were introduced to the New World
and their grazing and browsing also affected the forests,
frequently ensuring that regeneration did not occur after
clearing had taken place. It is, however, within the last 50
years or so that deforestation has really accelerated, and the
causes of this are broadly:
© to provide more land and wood (for fuel and building
materials) for a largely subsistence population,
particularly in countries with the greatest population
growth
to provide hard currency and vital export earnings from
the sale of timber, from cash crops grown on deforested
land and from the exploitation of minerals
the building of dams, roads, cities, etc. to meet the
needs of a growing urban population. These causes are
dealt with in more detail below.
Shifting cultivation
Shifting cultivation has been identified as the principal cause
of forest loss in all three tropical regions, accounting for
70% of the deforestation in Africa, 50% in Asia and 35%
in the Americas (Lanly, 1982).
Traditional shifting cultivators, however, who incorporate
forest fallow in their rotation, do not deforest. This method
involves clearing the forest and usually burning the wood so
that the nutrients within it are returned to the soil. Crops
will be planted for two or three years until the soil becomes
less fertile or weeds encroach on the area. The farmer then
moves on and repeats the process in another area, and only
returns to the initial patch after it has been left to regenerate
(left fallow) for at least 10 years and often more. With this
long fallow period to maintain soil fertility and in areas of
low population density (generally five or less per km?,
Myers, 1980), such systems are viable and the use of forest
land sustainable (Chin, 1987).
However, below a certain minimum fallow period
(depending on climatic factors, soil type, etc.) the forest
fallow develops a secondary thicket, yields drop and erosion
and soil degradation takes place. Forest removal becomes
265
permanent unless active attempts are made to reforest. This
permanent removal occurs where the numbers of shifting
cultivators have increased greatly; in Madagascar, for
instance, the population has risen from around 5 million in
1960 to nearly 12 million at present and tavy or shifting
cultivation is the main cause of forest loss in the moist
forests in the east of the country (see case study). In other
cases, not only are there too many people but these people
are immigrant farmers unused to local conditions who
frequently deplete the soil even more rapidly by using
inappropriate farming methods, and consequently clear ever
larger areas of forest. For example, in the south-west of
Céte dIvoire, smallholders successfully cultivate food crops
as well as coffee and cacao in a rain forest environment.
Their methods involve selective felling, thereby retaining
some forest trees, light burning, no tillage and maintaining
some fallow periods. In contrast, the immigrant farmers,
who have moved into the area from savanna lands, clear-
fell the forest, burn it heavily, and employ soil tillage
(Reitbergen, 1992).
Transmigration
Social and political factors often underlie forest loss. A
classic example is the transmigration programmes that have
occurred in Indonesia and Brazil (see case studies). A team
from the International Institute for Environment and
Development with representatives from three Indonesian
Ministries came to the conclusion that transmigration was
"the single sectoral activity with the greatest potential to
advance forest destruction - often to no constructive result.
The programme ... does not support the sustainable
development of Indonesia’s forest lands or, for that matter,
the settlements themselves" (Colchester, 1987).
In Brazil, deforestation by shifting cultivators, encouraged
to settle in the forests by the government, has been
compounded by large landowners moving in after them and
buying up the cleared land to convert to cattle ranches. The
immigrants then move on to clear more forest. Cattle
ranching in Latin America is estimated to have occupied
some 20,000km? of forest per year in the late 1970s
(Myers, 1990) and was profitable only because of the
1. Biological Diversity
some 20,000km? of forest per year in the late 1970s
(Myers, 1990) and was profitable only because of the
subsidies offered by the Brazilian government through tax
concessions and other such incentives. Once patches of the
forest have been felled, leaving piles of dry wood in an
area, the area becomes particularly susceptible to fires.
These have exacerbated the forest loss, particularly in the
Amazon where in 1987 some 50,000km? of forest were
burned in the two states of Rondonia and Acre alone
(Myers, 1990). Indeed, fires, either natural or deliberately
set, have caused forest destruction and prevented its
regeneration over huge areas, especially in Africa, for many
thousands of years.
Population pressure
The world’s population has risen from around three billion
people in 1960 to over five billion in 1990 and is estimated
to reach over eight billion by 2020. Although an inverse
relation appears to exist between population and forest
cover, the relationship is not precise (Mather, 1990). In
some countries, such as Kenya, the increase in numbers of
people has caused deforestation, but in many instances it is
the unequal distribution of land ownership rather than total
numbers that is the root cause of the problem. In countries
such as the Philippines, Brazil and Costa Rica (see case
study), most of the land is owned by a very small section
of the community. Most of the population has to derive a
living from very little land while elsewhere large areas are
underused or wrongly used. The pressure on the forests in
these instances stems from the inequality of the social
policies, compounded by giving priority to crops other than
those which ensure that the local population is fed
(Westoby, 1989). Indeed, Repetto (1990) suggests that
"government policies that encourage exploitation - in
particular excessive logging and clearing for ranches and
farms - are largely to blame for the accelerating destruction
of tropical forests."
Logging
Initially, most exploitation for tropical timber was for
highly valued species such as mahogany, teak and cedar. In
addition, until the 1940s the forests had been harvested
using axes, handsaws and animal power so that only forests
near rivers could be intensively exploited because of the
problem of extracting the logs. Under these circumstances,
the impact of logging forests was relatively minor.
However, the last few decades have seen a huge increase in
export of tropical timbers which owes much to the rising
affluence of the developed countries and the consequent rise
in the demand for hardwoods (Westoby, 1989). Advances
in technology, including the advent of chainsaws, tractors
and trucks and the buildings of roads and railways to reach
the most inaccessible areas have greatly facilitated the
exploitation of the forests. For example, the inland forests
of Gabon have, until recently, been protected because of
their inaccessibility but, with the building of the Trans-
Gabonese railway, the extraction and export of logs is
expected to rise considerably. Overall, exports of tropical
timbers to industrialised nations has risen sixteenfold since
1950 (Poore and Sayer, 1991). Information for Bangladesh
(Table 20.12) gives some indication of the increase in
extraction of timber between 1977 and 1984 as well as
bamboo (used for building houses) and firewood.
Inefficient logging practices contribute to the destruction of
the forest. For example, selection of mature trees of the
most valuable species may involve extraction of less than
10% of the timber in an area, yet it can typically result in
the destruction of at least half of the remaining stock,
including immature trees of the valued species as well as
harvestable stocks of somewhat less desirable trees
(Repetto, 1990). Repeated logging of partially harvested
areas to extract more timber before stands have recovered
can inflict heavy damage on the remaining trees and make
regeneration impossible (Repetto, 1990). In addition,
governments increasingly encourage local processing of the
timber, and in many cases the outdated machinery employed
in the local industry consumes considerably more timber
than efficient mills would expend to produce the same
output. This is the situation in Céte d’Ivoire where 30%
more logs are consumed by the local mills than would be
the case in an efficient operation (Repetto, 1990). In
Cameroon, the average conversion of sawn timber
processed locally for export is about 30% but can be as low
as 20% (Gartlan, 1989).
Roads and railways
One of the major indirect effects of logging is that the roads
built to enter an area are subsequently used by
agriculturalists to penetrate the forest, causing yet more
clearance. Indeed, in many instances the building of roads
and railways paves the way for further deforestation. For
Table 20.12 Output from forests in Bangladesh
OUTPUT (in thousand tonne) % CHANGE
ITEM
1977-78 1980-81 1983-84 1977-84
Timber 424 597 718 +69
Firewood 507 931 1279 +152
Bamboo 805 1449 1732 +115
Source: Ahmad, M. 1987. Bangladesh: how forest exploitation is leading to disaster. In: Forest Resource Crisis in the Third World. Sahabat Alam
Malaysia, Penang.
266
example, in Céte d’Ivoire it has been estimated that one
hectare of forest is removed by ‘follow-on’ cultivators for
every 5m? of timber removed by loggers (Myers, 1983). A
classic example of road construction attracting people to an
area and thereby accelerating deforestation is that of BR
364 in Rondonia, Brazil (see case study).
Large-scale commercial clearance
Conversion of forests to pasture is the foremost cause of
deforestation in the countries of Central America. In Brazil,
it has been estimated that 72% of the clearance in 1980 was
for pastureland (Browder, 1988). Forest continues to be
cleared on a large scale for plantations of oil palm, rubber,
sugar cane, tea, coffee, and cacao. In Peninsular Malaysia,
most clearance has been for agriculture, principally for oil
palm and rubber (Brookfield et al., 1990). The total area
under agriculture was 21% in 1966. This had risen to 39%
in 1982 and governmentconversion plans intend that around
45 % will be under agriculture by the mid-1990s (Brookfield
et al., 1990).
Fuel and charcoal
Domestic collections of firewood and charcoal are not
considered an important agent in the deforestation of the
moist forests, as most firewood is collected from savanna
woodlands, scrub and farmlands (Eckholm ef al., 1984;
Myers, 1980). However, the demand for fuel is rising
rapidly as populations increase and wood, both dead and
living, will inevitably be harvested in greater quantities
from the moist forests. In contrast, industrial consumers are
a cause of much outright deforestation (Eckholm ef al.
1984). For instance in Brazil, in addition to the forest
cleared to grow sugar, huge areas have been cut for the fuel
needed to process the cane (Williams, 1990).
Other factors
In itself, mining is a comparatively minor cause of
deforestation but the associated activities such as road
building and the discharge of chemicals and silt into rivers
can cause considerable damage. The same is true for oil
exploration, where it is the roads, and the pipelines and
spillage of oil and chemicals that are the principal causes of
the deforestation. War is an exceptional cause of
deforestation but, in the case of Viet Nam, bombing and the
extensive use of herbicides have destroyed large areas of
forest (see case study).
Summary
Overall, the causes of deforestation are many and varied,
and the impact of each differs between countries and even
between areas. There is widespread agreement that most
governments seriously underestimate the economic value of
their forests both as productive sources of commodities and
for the services they provide. The cost involved in
transforming the capital of the natural forest into other
forms of capital is not generally recognised. However, any
economic judgement of ‘the value’ of a forest must take
account of the long-term benefits of conservation, but
cannot ignore the short-term financial costs to tropical
countries which are already under severe economic
267
Tropical Moist Forests
constraints. Deforestation will be reversed only when the
natural forest is seen to be more economically valuable than
alternative uses for the land. Well-intentioned but misguided
policies by the developed countries, such as restrictions or
outright bans on the import of tropical hardwoods, may not
be in the best interests of forest conservation if such actions
reduce the immediate, direct value of the forests to the
exporting countries. Each country with tropical moist forest
within its sovereign control will have to develop, with the
support of the richer countries, its own particular repertoire
of methods to reverse the loss of its forests based on a
comprehensive understanding of the causal agents of that
loss.
MANAGEMENT PRACTICES
FORESTS
IN TROPICAL
There are a variety of management objectives for tropical
forests and these fall broadly into three categories: 1) for
the supply of products either timber or non-timber; 2) for
the protection of critical soils and water catchment; and 3)
for the conservation of biological diversity (Poore and
Sayer, 1991). The management practices applied to the
areas will obviously depend on the use of the forest but, in
a recent IUCN publication (Poore and Sayer, 1991) six key
principles for the management of tropical moist forest land
were listed.
Ecological constraints must be considered at the outset of
the development of tropical forest land.
The allocation of tropical forest land to other uses should
be decided only after thorough economic, social and
ecological evaluation, including consultation with local
communities
Tropical forest should be converted to uses other than
natural forest only if it can be demonstrated that this will
produce sustainable benefits in a more desirable form
than the original tropical forest itself can provide
Wherever possible, areas of tropical forest which are
already degraded should be selected for uses other than
natural forest rather than clearing pristine forest
Special efforts should be made to manage carefully those
large areas of tropical forest which are essential for
benefits such as the maintenance of watersheds and
biological diversity
The people who live in and around tropical forests
should have a major say in their management.
In essence then, the management of a tropical forest
involves the sustainable and continued harvest of all
products, including to satisfy the needs of the local people,
the maintenance of essential ecological processes, and the
conservation of biological diversity.
Management for biological diversity
The principal means of managing tropical forests for
biological diversity is through conservation areas. However,
only about 4% of the world’s remaining tropical forests are
legally protected, and in many cases these areas have no
management plans and no effective protection on the
ground. It is also unrealistic to anticipate expanding the
network of forest protected areas to cover all species and
ecological processes (Poore and Sayer, 1991). It is now
1. Biological Diversity
well appreciated that effective management must provide
real incentives, based upon an equable sharing of benefits,
for local people to participate in the conservation process.
Such benefits may be consumptive in the form of meat,
food, building materials, medicines and other forest
products, or non-consumptive, particularly in the form of
tourism.
Tourism is of increasing importance in the management of
forest protected areas, particularly for the income and local
employment it can provide. For example, tourists visiting
the Mountain Gorillas Gorilla gorilla berengei in the
Volcanoes National Park paid over US$800,000 in park fees
in 1989. With the distribution of these benefits amongst the
local community, together with increased educational
outreach, the proportion of local farmers who would like to
see the park degazetted has declined from 50% ten years
ago to only 20% (Harcourt et al., 1986). Incentives to local
communities in the form of increased access to credit,
capital grants, and support for improved agro-forestry
schemes are now increasingly used to encourage villages to
respect protected area boundaries.
Estimates of the extent of protected areas within tropical
moist forest are presented in Table 20.10. Very few
countries have more than 10% of their forest area under
protection, and only Burundi, Singapore, Australia and Sri
Lanka have over half their remaining forest under direct
conservation management.
Management for maintenance of service values
Most natural forests protect soil fertility, prevent soil
erosion, regulate water run-off and have a moderating effect
on climate. The purpose of protection forests is to maintain
these values. The services can also be provided to a lesser
extent by degraded forests or even artificial plantations, but
the maintenance of these ecological services is often
combined with preserving biodiversity values. Maintaining
natural protection forest also allows harvesting of non-
timber products, giving these areas both an immediate and
an indirect economic value.
Management for timber
Forests can be managed for timber at a number of different
levels of intensity:
e the lowest level is the demarcation of a remote area
which may be economically unattractive until shortages
of supply drive up market prices
an area may be selectively logged, protected from
encroachment to allow regeneration, and then
periodically relogged
selected trees may be extracted at a pre-set felling
intensity that ensures maximum economic return but
causes minimum damage to remaining trees that can be
harvested in the future
minimum intervention harvesting may be followed by
various. treatments, such as weeding or poisoning of
unwanted tree species
logging may be followed by enrichment planting of
saplings of economically desirable species.
In general, the more intense the management for timber
production, the less diverse the forest becomes.
At present, most of the supply of tropical hardwood timbers
comes from the first cut of previously unlogged forests.
Some timber does, however, come from forests that are
being relogged, mostly without a management plan, with a
small amount from areas that are being converted to
alternative use, mainly in an unplanned fashion.
Comparatively little comes from plantations, secondary
regrowth or agroforestry.
As the supply from the first cut of pristine forest and from
land destined for conversion to agriculture declines, the
immediate market reaction to the resultant shortages and
increase in price is to shift operations away from the
countries where supply has dropped to those which have
largely unused forest resources. For this reason, it is
predicted that there will be a movement away from
Southeast Asia, the major supplier at present, to South
America (Grainger, 1987). In all probability, these new
producer countries will then unsustainably mine their forests
in the same way as their predecessors have done.
A recent study undertaken by the International Tropical
Timber Organization (ITTO) (Poore et al. , 1989) found that
virtually nowhere was it possible to demonstrate
conclusively that any natural tropical forest had been
successfully managed for the sustainable production of
timber (see, for instance, case study on Congo). In Latin
America and the Caribbean, the total area being sustainably
managed at the operational level was limited to 75,000ha in
Trinidad and Tobago. In Australia, an area of some
160,000ha, the whole estate of tropical forest in Queensland
scheduled for logging was under sustainable production of
timber, although all this area has now been taken out of
logging following nomination as a World Heritage Site.
Of the total forested area of Asia, the only region practising
sustainable yield management was in parts of Peninsular
Malaysia where the ‘selective management system’ is in
operation. However, even here the system has only recently
come into full use and there is no certainty of its success
after the first cutting cycle, although it is intended to extend
the system to the total production forest estate of the
country. In Africa, the report concluded that there were no
sustained yield management systems currently being
practised throughout the continent. With the partial
exception of Ghana, forest management systems have been
progressively abandoned. A selection system similar to
Peninsular Malaysia has been running for eight years in the
Céte d’Ivoire, and preliminary results are sufficiently
encouraging for it to be extended to some 10,000ha of Yapo
Forest.
The report concludes that of an estimated total area of some
828 million ha of productive tropical forest remaining in
1985, the total under sustained yield managementamounted,
at the very most, to about one million ha. This is the reality
of the sophistication of tropical forest management
throughout the world. Urgent action is required not only to
ensure proper management of previously unlogged forests
but also to assess the status of logged forests and degraded
forest lands to bring these under sustainable protection. In
view of the scale of the task, the lack of an operational
definition of sustainability, and the need to develop
mechanisms to monitor forest management, this is an
ambitious target.
CASE STUDIES
Deforestation and degradation of forests in Sumatra
Figures for the original and present extent of rain forest in
Sumatra are not easily obtained, as sources such as FAO
(1988) give figures for Indonesia rather than for the
separate islands. However, a series of maps showing the
rapid disappearance of pristine forests in the country has
recently been published (Collins et al., 1991. See Fig.
20.1). In this same publication it is estimated that around
49% of the country’s land area is still forested but this
figure includes logged as well as the untouched forests
depicted in Fig. 20.1.
Figure 20.1 ‘Pristine forests in Sumatra
B&B Pristine forest
Source: Collins, N.M., Sayer, J.A. and Whitmore, T.C. (Eds) 1991.
The Conservation Atlas of Tropical Forests: Asia and the Pacific.
Macmillan Press, London, UK in collaboration with IUCN, Gland,
Switzerland.
The causes of the deforestation and degradation are varied,
but clearance for agricultural land is probably the primary
cause. Population density is high (59 people per km? in
269
Tropical Moist Forests
1980) and large areas have been cleared both for
subsistence agriculture and industrial plantations (Whitten
et al., 1984). In addition, relatively large areas of the
shallower peat swamp forests along the Malacca Straits
have been drained to provide farmland for settlers who
were moved there in the course of Indonesia’s large
transmigration scheme. There has also been considerable
logging in the country. For instance, on the flat lowlands of
southern Sumatra great stands of the commercially
important Ironwood Eusideroxylon zwageri, which produces
an exceptionally durable timber, have been almost entirely
destroyed while, in recent years, there has been heavy
logging in the lowlands east of the mountain spine. It is
probable that Sumatra is losing its natural vegetation faster
than any other part of Indonesia.
Deforestation in Viet Nam
An FAO (1987) report estimates that closed forest cover
was 61,650km? in 1980, 48,620 in 1985 and projected that
there would be only 34,060km? in 1990. In contrast, the
Ministry of Forestry (1989, unpublished) using 1987
Landsat imagery, indicated that 79,054km? of closed
broadleaved forest remained in 1987. Interpretation of what
is believed to be the same 1987 data set by WCMC gives
a total of 56,680km? of closed forest (MacKinnon and Cox,
1991). The variation in the statistics is, no doubt, due to
differing interpretations of what constitutes a closed canopy
forest.
The population of Viet Nam was originally centred on the
Red River Delta in the north but moved south during
historical times, clearing and cultivating the coastal plains
and valleys and reaching the Mekong Delta a few centuries
ago. These areas were consequently the first to be cleared
of forest. By 1943, about 45% of the country was still
forested. During the French colonial administration, which
ended in 1954, extensive areas in the south were further
cleared for industrial plantations, mostly banana, coffee and
rubber.
From 1945-1975, there was almost uninterrupted warfare in
the country. It was estimated that during the war between
the North and South, 22,000km?2 of forest and farmland
were destroyed by intensive bombing, spraying of herbicide
and mechanical clearing of forest. In total some 23,000-
55,000km? of forest were damaged by the bombardment
(Myers, 1980). In addition, large areas of forest were
cleared for agricultural land to feed the people (the
population doubled between 1945 and 1985). The area
cleared was larger than would normally have been needed
because of the necessity to make up for the food lost when
irrigation systems were bombed and crops were killed by
herbicides.
Since 1975, the need to rebuild after the war and the still
increasing growth in population has caused continuing loss
of forest. In 1981, FAO/UNEP estimated annual
deforestation of dense broadleaved forest during 1981-1985
at 600km? but FAO’s report of 1987 indicated the much
greater figure of 3,110km? of forest lost annually.
1. Biological Diversity
Figure 20.2 Viet Nam’s vanishing forest
cover 1943-1982
@ Forest cover
1975-1976
Source: Anon. 1985. Viet Nam: National Conservation Strategy.
Prepared by the Committee for Rational Utilization of Natural
Resources and Environmental Protection (Programme 52-02) with
assistance from IUCN. WWF-India, New Delhi.
Eastern rain forests of Madagascar
Estimates of forest extent and rate of deforestation in the
moist forests of Madagascar vary enormously. Myers
(1980), using reports from a variety of authors, estimated
that there were only 26,000km? of eastern rain forest
remaining in the country and that it was being deforested at
a rate of 2,000-3,000knr per year. In contrast, FAO/UNEP
(1981) reported over 69,000km? remaining in 1980 with an
annual deforestation rate for all the closed broadleaved
forest of 400km? between 1976 and 1980, which it was
predicted would decline during the period 1980-1985. In its
1988 report, FAO estimated an annual deforestation rate for
all closed broadleaved forests of 150km?.
A recent report, based on analysis of vegetation maps that
were made from aerial photographs in 1950 and on satellite
image data from 1984-1985, provides more accurate
estimates of remaining forest cover and deforestation rates
(Green and Sussman, 1990). The authors estimate that there
were originally 11.2 million ha of eastern rain forest, that
7.6 million ha remained in 1950, and that these had been
reduced to only 3.8 million ha by 1985 (Fig. 20.3). The
deforestation rate between 1950 and 1985 was, therefore,
111,000ha per year.
The main cause of the deforestation in the eastern rain
forests is slash-and-burn (or favy) agriculture, and cutting
for fuelwood to sustain Madagascar’s growing population.
The country supported 5.4 million people in 1960 and this
had risen to 12 million by 1990. The population is still
mostly rural and survives by subsistence agriculture. To
obtain more land, forest areas are clear cut, the vegetation
is dried and then fired some months later. Dry land rice is
most commonly planted, but maize, manioc and other crops
are also grown. They are cultivated for a year or two; then
the land is left fallow to regain its fertility and the process
is repeated elsewhere. Tavy has been practised for centuries
but the increase in population has put greater pressure on
270
Figure 20.3 Deforestation in
Madagascar’s eastern rain
forests
Source: Green, G.M. and Sussman, R.W. 1990. Deforestation history
of the eastern rain forests of Madagascar from satellite images. Science
248:212-215
the land, and it is now often left fallow for only three or
four years. As a result, the soil productivity progressively
deteriorates, and the area becomes unproductive grassland
or, on steep slopes, erodes away to bare earth. Predictably,
deforestation has been most rapid in areas of high
population density and low topographic relief. If cutting
continues at the present rate, Green and Sussman (1990)
estimate that only the forests on the steepest slopes will
survive the next 35 years.
Forest Loss in Costa Rica
Agricultural growth in Central America, as in other
developing countries, is driven by an expansion of pasture
and cropland rather than through intensified agriculture on
existing cleared land. The area under forest shrinks as a
result. The rate of transformation from forest to pasture has
been increasing since 1950. In Costa Rica, 67% of the
country was covered in primary forest in 1940, but only
17% remained under primary forest by 1983 and this was
mostly in the mountainous areas of the country (Fig. 20.4).
Deforestation has been greater in the dry western area than
in the mountains as the former is comparatively easy to
clear and maintain as pastureland by burning.
Figure 20.4
Loss of primary forest in
Costa Rica 1940-1983
HM Primary forest
Source: After Sader, S.A. and Joyce, A.T. 1988. Deforestation rates
and trends in Costa Rica 1940-1983. Biotropica 20(1):14.
In 1960, only 19% of the country was under permanent
pasture, while by 1980 this area had risen to 31% (FAO in
Leonard, 1987). Beef production takes up the majority of
the converted land, with 15,580km? being devoted to cattle
in 1980. This is in spite of the fact that the beef industry in
Central America is very inefficient, with levels of
productivity per hectare of land being considerably lower
than, for example, in the USA. As in the rest of Central
America, big ranchers in Costa Rica own most of the land:
landowners, with 60% of all farmers occupying only 4% of
the land.
Deforestation in Central America
Throughout Central America, the single most important
ecological change that is taking place as a result of the
current demographic pressures and economic trends is the
rapid and continuing conversion of forests to other land
uses (Leonard, 1987. Table 20.13). Almost all of Central
America was originally forested but it is estimated that now
only 40% of the seven countries are still forested (Fig.
20.5). Two-thirds of all the forest clearing has occurred
since 1950 and the rates of forest clearance have increased
in every decade since that date (Parson, 1976).
There are obviously some positive results arising from this
deforestation: for instance, the cattle ranching and farming,
both occurring on cleared forest land, are major generators
of employment, national income and export revenue in the
region. However, the economic contribution is
predominantly indirect, that is from the land cleared of the
timber rather than from the timber itself.
Despite the very rapid consumption of forests in recent
years, the timber industry is not a major force in most of
Central America. This is because much of the timber cut is
not harvested for commercial purposes; instead it is burned
in place or felled and not used. Only in Honduras does
commercial harvesting of timber contribute significantly to
the economy of the country. Even here it has been
estimated that forests with a commercial timber value of
$320 million are wasted each year (Leonard, 1987).
271
Tropical Moist Forests
Figure 20.5 Deforestation in Central
America 1950-1985
iss8s
a,
Hl Dense forest cover
Does not include coastal
mangrove forests and open
pine savanna
However, logging tracks do, here as elsewhere, open up the
forest to subsequent colonisation. Road building, which has
been a major goal of most governments in the region since
the 1960s, has the same effect (Leonard, 1987). Demand
for fuelwood is not, overall, a major force of deforestation
though in arid highland areas of Guatemala, Honduras and
El Salvador it does have an impact.
Deforestation in Peninsular Malaysia
In 1966, dryland forest on Peninsular Malaysia occupied
68% of the land area. It had declined to 54% by 1982 and
is now less than 50% (Brookfield and Byron, 1990) (Fig.
20.6). Swamp forest diminished from 14% to 10% of the
Peninsula in the same time, and now occupies about 8% of
the land. The major cause of this forest depletion has been
the conversion of land from forest to agriculture. Clearing
the forests for large-scale farming of cash crops began on
the west coast where rubber, coconut and then oil palm
plantations were developed. More recently, conversion of
forest has been undertaken by various federal and state
governments for land development schemes to provide
agricultural land and employment for landless families
moved from other parts of the country. Development plans
for the country encourage the further conversion of forest.
For instance, the Fourth Malaysia Plan (1981-1985)
suggested that another 6,075km? were to be cleared for
rubber plantations, 8,470km? for oil palm and 1,500km? for
settlements (Whitten, 1991).
1. Biological Diversity
Table 20.13 Status of lowland and tropical montane forests in Central America
COUNTRY REMAINING (1983)
PRIMARY FOREST
(km?)
Nicaragua 27,000
Guatemala 25,700
Panama 21,500
Honduras 19,300
Costa Rica 15,400
Belize 9,750
El Salvador fo)
Central America 118,650
CURRENT ANNUAL RATE % OF 1983
OF FOREST LOSS COVER LOST
(km?) ANNUALLY
1,000 3.7
600 2.3
500 2.3
700 3.6
600 3.9
32 0.3
o} =
3,432 2.9
Source: Nations, J.D. and Komer, D.I. 1983. Central America’s Tropical Rainforests: positive steps for survival. Ambio 12(5):232-238.
Figure 20.6 Forested areas of the Malay
Peninsula at various dates
SSE Forest
Hl Wetland forest
(7) Other land use
kilometres
Other, though comparatively minor, causes of forest loss
are construction of dams for irrigation, hydroelectric
schemes and mining, particularly for alluvial tin (Collins et
al, 1991). Logging, although rarely causing total
deforestation, does result in significant ecological damage
in some parts of the Peninsula (Fig. 20.7 shows remaining
areas of logged and unlogged forest). The logging roads
open up areas for cultivation and settlement. Shifting
cultivation by indigenous people is not an important cause
of deforestation in the region.
212
after
present
forest,
of
Figure 20.7. Remaining
completion
conversion plans
Largely undisturbed
upland forest
Largely disturbed/previously
harvested forest
Resettlement in Rondonia, Brazil
There have been various schemes in Brazil, backed by the
government and by outside agencies, to move people from
over-populated areas to the Amazon basin. An extensive
programme of road building has opened up the forest to
these settlement schemes and to other landless people
moving into the region of their own accord. For instance,
small farmers were expelled from the central-south of Brazil
by conversion of coffee and other labour intensive crops to
mechanised soybeans and wheat, and these people migrated
to the Amazonian frontier rather than to urban slums
(Fearnside, 1986).
Rondonia has one of the highest rates of deforestation in
Brazil’s Amazonia. In 1960 the state was uninhabited except
for some Amerindians and a few rubber gatherers but by
the late 1970s this region had as many as 5,000 people
moving in every month. Many of the immigrants were
landless people from the south of Brazil. In 1981, the
World Bank agreed to finance further development of the
area and a major road, BR-364, was paved which increased
movement into the State. The result has been progressive
deforestation which shows up clearly on satellite images of
the region (Fig. 20.8). The soils under the forest are
generally so poor that the settlers have to clear another area
within a year or two, or else sell their land to cattle
ranchers. Ranching used to be profitable, in spite of the
Figure 20.8 Deforestation in Rondonia,
Brazil
poor yields of either milk or meat, as a result of tax
incentives provided by the government, but these incentives
are no longer available.
This satellite imagery traces the progressive clearance
(shown in black) of forest for farmland in an area of
Rondonia, south-west Amazonia. The vertical lines are
roads set Skm apart while the thicker black, curved line is
BR-364.
Transmigration as cause of deforestation in Indonesia
In Indonesia, nearly three million people have been moved
from the crowded and environmentally degraded islands of
Lombak, Bali, Java and Madura to new settlements on less
populated islands (Whitten, 1991). This has been the
world’s largest programme of voluntary assisted migration
but, in addition to the sponsored migrants, it is estimated
that two or three times as many unassisted people have
moved to the less populated islands.
The land to which the transmigrants have been moved is, in
many cases, entirely inappropriate. Some of the sites are
intrinsically unsuitable for agriculture; others were
inadequately prepared or inappropriately managed. As a
result, loss of forests throughout the region is one of the
major environmental impacts of the settlement programmes.
In addition, in many cases, the removal of people from
degraded land has not improved the environmental
273
Tropical Moist Forests
conditions they left behind. For example, in Java the
population has been growing faster than the rate of
transmigration and there does not appear to be any
improvement in the state of the critically eroded land on
this island.
The budget for this programme has been cut recently and it
has been agreed that no new areas are to be cleared.
Instead, development of the already existing sites will take
place, roads will be built to improve communications, tree
crops will be encouraged, produce will be marketed more
effectively, and other improvements will be made.
Fires in Borneo
Until recently, fires were not considered to be a major
factor in the fate of tropical forests. However, in 1982-
1983, major fires occurred in Borneo during a drought and
very large areas of forested land were burnt. Beginning late
in 1982 and peaking in early 1983, numerous fires broke
out in coastal and inland areas of East Kalimantan. In
Sabah, an overlapping series of outbreaks occurred from
early through mid-1983. Minor outbreaks also occurred in
other parts of Borneo and the southern Peninsula.
The lowlands of East Kalimantan are one of the driest areas
of the island of Borneo, and during the intense El Nino
southern oscillation of 1982-1983 rainfall was reduced by
more than 60% compared with long-term monthly averages.
From February to May 1983, instead of receiving more
than 135mm rain per month as normal, almost none fell
(Malingreau et al., 1985). Drought stress led to the
shedding of leaves by evergreen species and to the
accumulation of dry litter on the forest floor. The extensive
fires that took place, particularly from August to October
1982 and March to May 1983, were mainly triggered by the
agricultural practices used in the area, which included dry
season burning as a land clearing method (Malingreau et
al., 1985). Accelerated settlement programmes and
spontaneous migration have meant that large tracts of land
in East Kalimantan are being deforested for agricultural
uses, especially along the coast and main rivers, and
burning in these areas almost certainly caused the major
fires in that region.
It has been estimated that in East Kalimantan alone
35,000km? of land have been damaged by the fires. This
includes 8,000km? of primary lowland forest, 5,500km? of
peat swamp forest, 12,000km* of selectively logged forest
and 7,500km? of shifting cultivation land (Leighton and
Wirawan, 1986). It is thought that around 10,000km? of
vegetation were damaged in Sabah (Malingreau et al.,
1985). The data suggest that selectively logged forest
suffered greater damage than the primary forest, as the
debris resulting from the logging provided further dry
material to fuel the fires.
In conclusion, it is evident that increasing populations, with
the resulting increase in slash-and-burn agriculture,
combined with the extreme climatic conditions of 1982-1983
were the cause of considerable deforestation in Borneo. It
appears that fires may well become a more common agent
of deforestation in the future. Indeed, at the time of writing,
considerable areas of forest on the island are once more
ablaze.
1. Biological Diversity
in Borneo,
Figure 20.9 Forest fires
1982-1983
BORNEO
KALIMANTAN
Areas affected
by fires
&
14E 300 km
Forest management in Congo
Congo’s national forest estate has been divided into forest
management units, each of sufficient size to support an
independent forest industry. The industries are required to
conduct an inventory of their unit and propose a
management plan for ministerial approval. These plans
should provide for selection felling on a 25 year cycle with
a minimum diameter limit of 60cm. Extraction is subject to
three year exploitation permits, which prescribe the
maximum area to be logged and the minimum volume of
timber to be produced. This system could have provided a
sound basis for a sustainable forest industry but a variety of
factors have prevented it from ever being put into practice
properly.
Understaffing of the forestry service has meant that the
units are not properly supervised. More important, all forest
land is state property so even if an area is under
management for timber, all citizens have rights to use the
area. These customary rights not only allow subsistence
hunting and collecting of non-timber products (neither of
which harm the forest to any great extent), they also allow
local people to practise shifting cultivation in the area. This
has happened in the more densely populated south of the
country and potential timber yields have been significantly
reduced as a result. It appears that sustainability in the
south can be achieved only in intensively managed
plantations taken out of state ownership. In contrast, in the
comparatively inaccessible, sparsely populated north, the
forests remain undisturbed after logging and regenerate
well. Here selective logging is practised and a near natural
forest is maintained.
| ee)
274
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Myers, N. 1988. Tropical deforestation and remote sensing. Forest
Ecology and Management 23:215-225.
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Climatic Implications. Friends of the Earth, London.
Myers, N. 1990. The world’s forests and human populations: the
environmental interconnections. Population and Development
Review No.16.
Nations, J.D. and Komer, D.I. 1983. Central America’s Tropical
Rainforests: positive steps for survival. Ambio 12(5):232-238.
Parson, J.J. 1976. Forest to Pasture: development or destruction?
Revista de Biologia Tropical 24:121-138.
Peters, C.M., Gentry, A.H. and Mendelsohn, R.O. 1989. Valuation
of an Amazonian rainforest. Nature 339:29.
Philippines Forest Management Bureau, Department of Environment
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Philippines.
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1989. No Timber Without Trees. Earthscan, London.
Poore, D. and Sayer, J. 1991. The Management of Tropical Moist
Forest Lands: ecological guidelines, 2nd edn. TUCN, Gland,
Switzerland and Cambridge, UK. 76pp.
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C.S. and Collins, N.M. (Eds), The Conservation Atlas of Tropical
Forests: Africa. Macmillian Press, London, UK in collaboration
with IUCN, Gland, Switzerland.
275
Tropical Moist Forests
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262(4):18-24.
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London, UK in collaboration with IUCN, Gland, Switzerland.
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destruction and species extinction. Biological Conservation
55:199-213.
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Chapter written by Caroline Harcourt. We thank
Biotropica, H. Brookfield and Y. Byron, P.M. Fearnside,
G. Stephens, and R.W. Sussman for giving permission to
reproduce maps originally published by them.
1. Biological Diversity
Figure 20.10 Tropical Moist Forest: Asia
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276
Tropical Moist Forests
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277
1. Biological Diversity
Figure 20.12 Tropical Moist Forest:
Africa
278
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279
1. Biological Diversity
21. GRASSLANDS
Grassland can be described as a type of vegetation that is
subjected to periodic drought, is dominated by grass and
grass-like species, and grows where there are fewer than
10-15 trees per hectare. This definition is somewhat
arbitrary and is one of several that may be used in
discussion of grasslands and the area they cover. Different
vernacular terms are used depending on the part of the
world under consideration; thus grasslands may be called
steppes in Eurasia, prairies in North America, llanos,
cerrados or pampas in South America, savannas in Africa
and rangelands in Australia.
Although virtually all the world’s grasslands have been
affected to some extent by man and his domestic stock,
natural grasslands appear to have developed in two kinds of
area:
areas where the growth of trees is prevented by edaphic
or climatic factors (these are limited in extent, being
confined to areas with nutrient-poor soil and / or low
rainfall)
areas where, over a very long period, browsing by wild
herbivores has prevented the establishment and growth
of trees, and where wild species are still present and
outnumber domestic livestock.
In both these types of natural grassland the dominant
ecological factors have prevailed for sufficiently long for
plants and animals to have adapted and established a natural
balance. Two of the essential qualities of such natural
grasslands are that the vegetation is unsown and that the
balance between plant species has not been significantly
affected by human activity. From these natural grasslands
there is then a complete spectrum of degrees of
modification by man, finishing with the entirely sown and
intensively managed short-term rye-grass ley of western
Europe, which has almost no significance for the
maintenance of biodiversity. Semi-natural grasslands which
are unsown but strongly modified by the grazing of
domestic livestock are of much greater importance. A large
proportion of the world’s grassland species are able to use
such habitats, and many species are, indeed, dependent on
them.
Many natural and semi-natural grasslands have high levels
of floristic diversity, at some scales and in some areas
approaching that of tropical forests. Animal species richness
appears to be generally low, although only vertebrates are
well recorded. For example, those birds that are considered
to be primarily adapted to grasslands and dependent on
them number around 477 species worldwide; this is less
than 5% of the world’s bird species. One characteristic of
grassland birds is a tendency for rapid and apparently
erratic dispersal, which enables them to exploit sparsely
distributed food resources in an environment in which the
climate is unpredictable. Many species habitually move over
very large areas. These factors mean that it is difficult to
conserve grassland birds through protection and
management of important wildlife areas (Grimmett and
Jones, 1989). Many more species would by now have
become globally threatened were it not for their ability to
use mixed farmland.
280
Similarly, a total of 245 of the world’s mammal species are
considered to be primarily adapted to grassland conditions.
This represents about 6% of the world’s described mammal
species. They can be broadly divided into large and small
mammals, with 77 of the former and 168 of the latter. The
overwhelming majority of the large mammals are grazers
and/or browsers, and only 19 are predators or scavengers.
In general, mammalian predators are adapted to a range of
habitats rather than being restricted to grassland. The small
mammals of grasslands are mostly seed-eaters or
omnivores.
THE WORLD AREA OF GRASSLAND
It has been estimated that grasslands covered approximately
40% of the earth’s surface prior to the impact of man and
his domesticated animals (Clements and Shelford, 1939).
Estimates of the area of grassland present today are
generally much lower than this but are very variable. One
of the highest estimates suggests that grasslands occupy
27% of the world’s natural vegetation cover (Knystautas,
1987). Data on savanna and temperate grasslands from
other sources are incorporated here into Table 21.1 and the
percentage of the world’s land area occupied by these
habitats has been calculated. These result in estimates of the
area of savanna and temperate grasslands ranging from
16.1% to 23.7% of the world’s land area (or 17.9% to
26.5% if Antarctica is excluded).
The land-use statistics produced by FAO (e.g. FAO, 1987)
on a country basis include a category for pasture, but this
is not clearly defined and it is certainly not restricted to
long established, semi-natural or biologically significant
habitats. A great deal of the pasture is either newly created
from woodland or arable, and it may be managed in
rotation with other farm crops. Other areas regarded as
permanent pasture by FAO may have had long continuity of
grazing but their native flora and fauna may have been
completely lost because of agricultural intensification.
Whatever the actual area of grassland present today, two
facts are clear: there used to be considerably more natural
grassland in the world, and its area is continuing to
diminish.
ORIGINS AND FLORAL DIVERSITY OF
GRASSLANDS
The best examples of grasslands that formed where soil and
climate favoured the production of grass and herbaceous
species, rather than trees, are those in northern South
America and in South Africa.
In other parts of the world, the impact of large herbivores
was more significant. The main areas where grassland
formation was influenced by large herbivores were the
savanna zones of Africa, the steppes of Asia and Eastern
Europe, and the prairies of North America. In Africa the
large wild herbivore community was dominated by
ungulates such as antelopes and zebras; in Eurasia by
gazelles, goats, camels, bison and wild horses; and in North
America by deer and the North American bison. The effects
Grasslands
Table 21.1. +Estimates of the area* of the world’s grasslands
WHITTAKER ATLAY, KETNER OLSON, WATTS
AND LIKENS AND DUVIGNEAUD AND ALLISON
(1975) (1979) (1983)
Savanna 15.0 22.5 24.6
Temperate grassland 9.0 12.5 6.7
Total grassland 24.0 35.0 SieS
Grassland as % of world land area 16.1% 23.7% 20.7%
Grassland as % of world land area (excluding
Antarctica) 17.9% 26.5% 23.1%
Note: * in million km?.
of these larger species were supplemented by vast numbers
of small mammals such as marmots, pikas, ground
squirrels, gerbils and voles. In the African savanna and
Australian rangelands, termites are extremely important:
they may consume up to one-third of the total annual
production of dead wood, leaves and grass, and their
biomass may reach as high as 22g/m’, more than twice that
of the greatest densities of vertebrates on Earth, found in
the migrating herds of ungulates on the Serengeti plains,
Tanzania.
South America
Long-established, near-natural savannas occur in tropical
South America in regions where the climate is in no way
inimical to the growth of trees. These tropical savannas
occur over huge areas as a mosaic of grassland and forest,
sometimes as extensive grassy plains with scattered trees,
sometimes as grassland with strips of woodland and
sometimes as islands of grassland in vast tracts of forest.
Fig. 21.1 from Sarmiento (1983) shows the distribution of
Figure 21.1 South America: major
tropical savanna regions
fq Tropical
Esq] Trop!
Savanna
1000
Source:
Savannas. Ecosystems of the World, 13. Elsevier, Amsterdam.
Sarmiento, G. 1983. In: Boulitre, F. (Ed.), Tropical
281
these habitats. To the south of the area shown, the
grasslands are all secondary and less biologically diverse.
The main areas of savanna are listed in Table 21.2, along
with their area and floristic richness. The numbers of
species refer to entire regions, so they include species
which are not primarily adapted to grasslands. It is difficult
to make comparisons between the regions but two areas
stand out as being both large in extent and rich in plant
species other than trees and shrubs. These are the
Colombian-Venezuelan //anos, to the west and north of the
Orinoco river, and the very extensive central Brazilian
cerrados. Both are rich in plant species and communities,
the difference between them being related mainly to soils
and drainage (Huber, 1987).
Sarmiento gives a few figures for floristic richness of the
cerrados on a smaller, repeatable scale but unfortunately
comparable data from other areas have not been located. He
states that more than 300 plant species are recorded per
hectare of protected cerrados near Brasilia. The sampling
was done in 20 x 20 metre plots, with the number of
species per plot varying from 52 to 117.
It is interesting to note that although the numbers of species
in these neotropical savanna formations (Table 21.2) are
generally high, they are not as rich in grasses and herbs as
some of the temperate South American grasslands. The
pampas of Argentina and Uruguay, which is of secondary
origin, has over 400 species of grasses (Cabrera, 1970).
The pampas has generally been grazed by cattle throughout
historic times. This has encouraged colonisation by species
which would not otherwise be able to compete with more
aggressive species. The pampas is, therefore, more
species-rich but less natural than the cerrados.
Fire plays a more important role than herbivores in
maintaining these South American savannas. Fire is a
naturally occurring phenomenon but man has increased the
frequency of burning in recent centuries and this has had a
significant influence on plant communities. Nutrient
deficiency, usually related to aluminium toxicity and water
availability (Folster and Huber, 1984), is also believed to be
of critical importance in maintaining the openness of the
vegetation. The largest of the secondary grasslands are the
pampas and the campos. The latter are open rolling plains
on the central plateau of interior Brazil. The campos tend
to merge into the cerrados, with a gradual increase in the
proportion of trees. All these areas have been extensively
modified by frequent burning and agricultural development.
1. Biological Diversity
Table 21.2 __Floristic richness of various Neotropical savanna formations
FORMATION AREA NO. OF NO. OF NO. TOTAL
(km?) TREES SUBSHRUBS OF NO.
AND SHRUBS HALF-SHRUBS GRASS OF
HERBS SPECIES SPECIES
VINES, etc
Cerrado in north-western 50 45 175 17 237
Sao Paulo
Cerrado in western Minas 15,000 c. 200 c. 330 73 c. 600
Gerais
Whole cerrado region 2,000,000 429 (774)! 181 108 718 (1063)'
Rio Branco savannas 40,000 40 87 9 136
Rupununi savannas 12,000 c. 50 291 90 431
Northern Suriname savannas c. 3,000 15 213 44 272 (445)?
Central Venezuelan Ilanos 3 69 (16)8 175 44 288
Venezuelan llanos 250,000 43 312 200 555
Colombian llanos 150,000 44 174 88 306
Notes: ' Total flora including other plant formations; * Total flora including bushes; * Number of savanna trees excluding groves. See Sarmiento,
1983, for data sources.
Africa
The parts of Africa covered by the various forms of
vegetation described as savanna are shown in Fig. 21.2,
reproduced from Menaut (1983). Everything which is not
forest, desert or montane vegetation is regarded as a form
of savanna, though clearly not all of this is grassland. Much
of the land is cultivated and probably an even bigger area
is savanna woodland. Nevertheless, a high proportion of
Africa supports dry, semi-natural vegetation in vast
unbroken tracts. The considerable age of the habitat and its
great geographical continuity are vital factors in explaining
the biodiversity of African grasslands.
Table 21.3 Africa: areal richness zones
REGION RICHNESS
Guineo-Congolese region, peripheral domain
Northern district 1,440
Southern district 1,680
Sudano-Zambezian region
Sahelian and Sudanian domains 1,060
Zambezian domain 2,590
Eastern transition zone
Sahelian type 1,270
Sudano-Zambezian type 2,330
Kalahari domain 1,020
Madagascar 5,410
Source: Menaut, J.-C., 1983. The vegetation of African savannas. In:
Boulitre, F. (Ed.), Tropical Savannas. Ecosystems of the World, 13.
Elsevier, Amsterdam.
The best information available on floristic richness of dry
tropical Africa was assembled by Lebrun and summarised
by Menaut (1983). Lebrun pointed out that to compare the
species richness of different areas it was essential that the
comparisons were made between units of similar size. He
chose a unit of 10,000km* as a standard and called the
average number of species in that area the areal richness of
the region. Table 21.3 shows the average areal richness for
the major plant-geographic (phytogeographic or
282
chorological) zones of Africa and Fig. 21.3 shows their
location. The data in Table 21.3 should not be used in a
detailed comparison of the diversity of grasslands because
a high proportion of the plant species listed are associated
with forest, wetland or other habitats. Nevertheless, the
zones show a high degree of correlation with the main
savanna zones, so some broad comparisons are possible.
Menaut points out that in Africa the average areal richness
for savanna (c. 1,750 species) is not far below that of rain
forest (c. 2,020 species), contrary to the situation in the
Americas. Indeed, the heart of the savanna zone, known as
the Somali-Masai Region, contains 2,500 plant species, of
which 50% are ecological endemics (Stuart and Adams,
1990). Not all of these can be regarded as primarily adapted
to grasslands, but a sufficiently large number are for it to
be regarded as the world’s richest grassland zone.
Biodiversity in the region is enhanced by the fact that the
savanna merges gradually into other large habitat
formations, notably forest and semi-desert, rather than
being confined by mountains, the sea or intensive
agriculture.
The presence of large mammals is important in the ecology
of African grasslands; the world’s greatest concentration of
large mammals is found on the savanna of northern
Tanzania. However, as in South America, fire has also been
a major influence in the evolution of the flora and fauna.
Natural fires, caused by lightning, may have affected huge
areas, limiting the build-up of dry organic matter and
favouring the survival of some species at the expense of
others. In addition, at least in East Africa, man has
probably been burning grasslands for a minimum of 50,000
years, and very likely much longer. Therefore, here as in
South America, man has greatly affected the apparently
‘natural’ grasslands.
Australia
The grasslands of Australia have been described by Moore
(1970) using a very broad definition of grasslands which
includes all regions where grasses supply a substantial
proportion of the food for stock. This includes a wide range
Figure 21.2 Africa main savanna
vegetation types
ASg 2
SS 2
Source: Menaut, J.-C., 1983. The vegetation of African savannas. In:
Bouliére, F. (Ed.), Tropical Savannas. Ecosystems of the World, 13.
Elsevier, Amsterdam.
Notes: 1 woodland; 2 tree/shrub savanna; 3 forest/savanna mosaic; 4
tree/shrub ’steppes’.
Figure 21.3 Africa: areal richness zones
[_) 9-500
[J soo-1000
1000-1500 "SC ee EDS
(SN) 1500-2000
KS§ 2000-3000
HI >s000
Source: Menaut, J.-C., 1983. The vegetation of African savannas. In:
Bouliére, F. (Ed.), Tropical Savannas. Ecosystems of the World, 13.
Elsevier, Amsterdam.
Notes: Areal richness measured by number of species per 10,000km’.
of semi-desert, scrub and wooded savanna. Groves (1981)
considers that the only natural grasslands in Australia are
283
Grasslands
those dominated by hummock grasses, i.e. species of
Triodia and Plectrachne. These are very similar to the
bunch grasses of North America. Hummock grassland is
distributed over a very large area of south, west and
northern Australia, in arid and semi-arid lands, but a high
proportion of this vegetation has been subject to agricultural
improvement. Relatively small areas are free of introduced
species, legumes being the most common of these. Legumes
raise soil nutrient levels and fertility, thus changing the
ecological balance and making conditions more suitable for
weedy species. Around the great zone of natural hummock
grasslands are other natural and semi-natural communities,
most of which contain a significant proportion of grasses.
The tropical zone of northern Queensland is strongly
influenced by summer rains, whereas temperate and less
natural grasslands occur in a zone from north of Adelaide
to northern New South Wales.
One of the main reasons for the difference in the vegetation
of the arid zone of Australia and, for instance, East Africa
was the limited number of indigenous grazing mammals in
Australia. The larger marsupials (wallabies and kangaroos)
are primarily browsers rather than grazers, and would have
had little impact on the arid hummock grassland. The
influence of herbivores was therefore of much less
significance than in other parts of the world. However,
traditional burning patterns used by the aborigines (who
arrived around 40,000 years ago), together with their
hunting practices, must have influenced plant communities
by favouring species adapted to fire.
The arrival in Australia of settlers from Europe and their
domestic animals, particularly sheep, led to the
establishment of very extensive rangelands in which grasses
were able to provide most of the fodder for stock. Many of
the indigenous species were able to adapt to these new
conditions but numerous plant and animal species were
introduced from other parts of the world also. The most
significant of these was the rabbit Oryctolagus cuniculus,
whose numbers exploded in the absence of other competing
herbivores. Australia’s grassland species are therefore a
complex mixture of desert- and fire-adapted species, species
secondarily adapted to grassland and introductions.
Asia
The natural Asiatic steppe extended from Manchuria
westwards as far as the land now occupied by Bulgaria and
Hungary, occupying the broad zone between the taiga
(coniferous boreal forest) and the deserts or mountains to
the south. The continental climate of this vast area, with
hot, dry summers and very cold winters, is inimical to the
growth of trees. A large proportion of the area has not
supported forest since a more favourable climate prevailed
in one of the earlier interglacials. The dominant herbivores
before the influence of man became widespread were
horses, wild sheep and gazelles, along with a wide range of
smaller mammals.
A large proportion of the Indian sub-continent supports
either tropical savanna, savanna woodland or dry forest.
The total range of grassland types is very broad - from
semi-desert, to seasonally inundated areas, to montane
habitats. In the case of India, there is no doubt that a very
1. Biological Diversity
large area formerly supported dry woodland of various
sorts, but there has been an enormous amount of clearance,
followed by fire and grazing. It appears likely that there are
no surviving primary grasslands in the country (although
there is continuing dispute over the origin of hill grasslands
in the south-west). In addition, there are only a few
long-established stable communities in which the balance of
species clearly reflects edaphic factors and traditional
management. Indian grasslands are apparently more seral in
character, always being in a phase of recovery from
clearance, fire, overgrazing, erosion or abandonment. This
exerts a powerful influence on the flora and fauna present.
The vegetation is relatively poorly endowed with perennial
herbaceous plants and floristic diversity is not particularly
high. Mammals are not well-represented but a large range
of bird species are able to use the grasslands and many of
them are dependent on it.
Europe
The Asiatic steppes were extended at an early stage by
forest clearance, initially to increase the grazing for sheep,
goats and horses. Islands of forest on better soils or where
rainfall was higher, were reduced or eliminated. The
clearance continued westwards into Europe, into regions
where the climate and soils were far more suited to the
growth of deciduous forest. Thus wholly new types of
grassland were created, capable of far greater productivity
than the natural steppes. Man’s activities enabled numerous
grassland plants, particularly shorter grasses and herbs, to
extend their range, accompanied by characteristic animals.
The extension of grazing enabled semi-natural grasslands to
develop westwards, as far as central Spain, the Atlantic
regions of France and the British Isles. While many species
expanded their range from the steppes, others were no
doubt lost altogether. It is very doubtful if any of the
European steppes can be regarded as primary (Polunin and
Walters, 1985) but some areas of secondary steppe may be
very similar to the original habitats. The most natural
grasslands in Europe are the Hungarian pusztas,
traditionally managed, low-lying grasslands in the floodplain
of the River Danube.
Man started grazing domestic stock on the mountain ranges
of Europe at a very early time and permanent settlements
were established high in the valleys. Extensive grazing by
sheep and goats occurred over the high mountains in the
summer but the stock was returned to pastures near the
farms for the winter. Much of the land on the lower slopes
was cut for hay to provide fodder for the long winters. The
pattern of management was so consistent that many different
plants were able to adapt to these conditions and the
meadows became very species-rich. The flora of alpine hay
meadows are a mixture of steppe and montane species.
They form a balanced semi-natural community reflecting
traditional patterns of land-use and are highly valued both
aesthetically and scientifically. It must be remembered,
however, that their origins are not natural.
The British Isles is a region lacking natural grasslands
(according to the definition above). Here too, however, the
pattern of pastoral land-use is sufficiently long-established
and consistent for man-made grasslands to appear natural
284
and they vary in composition in relation to soils, aspect and
drainage. As a result, these habitats have assumed great
value as resources for a variety of wildlife. For example, at
least a third of Britain’s 1,500 species of flowering plant
are associated with grassland, and about 400 of them are
most frequent in this habitat (Duffey et al., 1974). In
southern England, agriculturally unimproved, semi-natural
grasslands may have up to 40 higher plant species per m’.
Even this level of species richness is exceeded by natural
grasslands in the heart of the steppe region, which in parts
of the former USSR and Mongolia may support up to 80
species per m* (Knystautas, 1987).
North America
In the central parts of North America, in the rain shadow
of the Rockies, the dry climate naturally favours open
habitats rather than woodland. Vast herds of bison also
helped maintain species-rich grassland on a grand scale.
Islands
The absence of large wild grazing mammals from oceanic
islands ensures that they do not generally support natural
grasslands. Even where man has created grasslands on
islands, such as on Madagascar or Sri Lanka, they seldom
support diverse stable communities. The average areal
richness of Madagascar is higher than all the other
chorological territories in Africa, but this is because of the
very large number of forest species. Savannas have been
created in the drier parts of the island but they are
species-poor, with a high proportion of introduced plants
(Menaut, 1983).
An exception to this rule is New Zealand, which has some
long established grassland’ habitats. These are however
high altitude, high rainfall, tussocky communities, which
are very different from the rest of the world’s grasslands.
They support distinctive native species such as the Takahe
Porphyrio mantelli, a flightless bird of the family Rallidae.
Those grasslands which have been created in New Zealand
for grazing stock are very largely composed of introduced
plant species, incidentally supporting introduced wild
animals such as the Red Deer Cervus elaphus, and they
should be regarded as artificial.
The only island lacking large indigenous grazing herbivores
but supporting species-rich grassland is Cuba. Floristically,
the Cuban savannas compare favourably with their
equivalents in South America. This may be because of
climatic stability and greater ecological diversification on
several different types of parent material, including some
unusual substrates such as silicious rocks and serpentine
(Sarmiento, 1983). It seems very surprising, however, that
such diversity of grassland species could evolve in the
absence of large herbivores.
THE 20TH CENTURY IMPACT ON GRASSLANDS
Until this century, the distribution of grassland species
around the world had been determined by an integrated
complex of various factors, including: climate, geographic
and ecological isolation, the impact of large herbivores,
traditional land-use practices, domestication of grazing
animals and forest clearance. The richest grassland regions
of the world, in descending order of importance for
indigenous plants and animals, were:African savanna;
Eurasian steppe; South American savanna; North American
prairies; Indian savanna; and Australian grasslands.
On one hand, the original extent of these natural grassy
areas has been extended by man’s activities so that
species-rich, semi-natural grassland now occurs in a
discontinuous manner over a very much larger area. It is
found throughout much of the region once occupied by the
world’s temperate forests and also reaches well into the
tropical forest zone. On the other hand, domestic stock has
often overgrazed natural grasslands, causing massive
impoverishment of the ecosystem, and large areas have
been converted to agricultural land.
Africa
In Africa, native people have burnt the savanna for
thousands of years to improve grazing for their stock and
facilitate the hunting of wild game. The frequency of these
fires may have increased markedly over the last thousand
years or so as population increased. Towards the end of the
19th century, when settlement became more firmly
established, the area of savanna was greatly enlarged by
forest clearance, burning, and massive increases in the
number of cattle. Much of this was achieved by white
settlers with imported European stock. Their farming
activities were, however, frequently upset by the
unpredictability of the climate and by parasites and diseases
which became increasingly serious as livestock densities
rose. Many of the imported breeds proved incapable of
tolerating the indigenous diseases of Africa, the most
serious of which were rinderpest, trypanosomiasis and foot-
and-mouth disease.
Around the turn of the century, rinderpest spread from the
north of Africa to the far south in a period of only seven
years, killing 90-95% of domestic cattle as well as many
wild ungulates (Rogers and Randolph, 1988). The result
was widespread human starvation and the abandonment of
vast areas of grazing land. The subsequent regeneration of
scrub and woodland appears to have allowed unprecedented
spread of tsetse flies (vectors for the parasite causing
sleeping sickness in humans and trypanosomiasis in cattle).
Colonial governments tried a variety of methods in an
attempt to eradicate the tsetse fly, one of which was the
removal of the woodland and cover which the flies require.
This allowed the indigenous herbivores and, consequently,
their predators to extend their ranges. These enlarged
distributions have generally been maintained, assisted in
recent decades by control of poaching and the establishment
of national parks and other protected areas. The grassland
habitat in these areas may have every appearance of
naturalness but frequently it is not very old and its extent
has been greatly influenced by man.
There have been large increases in the area used for
growing cotton in some of the semi-arid parts of Africa,
particularly Senegal, Mali and Mauritania. Persistent
insecticides such as dieldrin have been used on these crops,
with little regard for non-target species. The crops provide
very little food for birds or mammals and the water control
285
Grasslands
schemes required for their irrigation intercepts water which
formerly flooded river valleys, where it provided suitable
feeding areas for many species, including migrant birds
(Goriup and Schulz, 1991). The loss of grassland presents
a particularly serious problem for European breeding birds
which winter in West Africa because the habitat is confined
on its southern edge by forest, farmland or the Atlantic
Ocean. In East Africa, on the other hand, there are few
barriers to prevent wintering birds from moving further
south to find suitable habitat.
Europe
A useful review of the surviving area of grassland in
Europe has been carried out by van Dijk (1991), and a
summary of this data is presented in Table 21.4. The
figures quoted for permanent pasture generally correspond
well with the latest FAO data. The majority of the
discrepancy in total area between the two datasets is
attributable to the UK although there are also major
differences in the figures for Greece and Spain. In the UK
the difference arises because upland areas in Scotland and
Northern Ireland were included in the FAO data but not in
van Dijk’s data. Wherever possible van Dijk used national
surveys (of various dates) but for other countries he has
drawn the information from Lee (1990) or Grimmett and
Jones (1989).
land-use 1970-
Figure 21.4 France:
1985
Source: Lecomte, P. and Voisin, S. 1991. Dry grassland birds in
France: status, distribution and conservation measures. In: Goriup,
P.D., Batten, L. and Norton, J. (Eds), The Conservation of Lowland
Dry Grassland Birds in Europe. Proceedings of an International
Seminar held at the University of Reading 20-22 March 1991.
The data clearly reveal that only a small proportion of
permanent grassland can still be regarded as dry
semi-natural, and therefore of biological interest. The
review by van Dijk gives a lot of information on habitat
loss in different countries and it is apparent from this that
a great deal of the loss has taken place since the 1960s.
More information on these trends is provided in the case
study on grasslands in Poland.
1. Biological Diversity
Table 21.4 Areas of grassland and dry semi-natural grassland in Europe (1,000ha)
COUNTRY FAO PERM VAN DIJK DRY SN PER CENT
(A) (B) (C) (C/B)
Belgium 688 632 0.5 + <1
Czechoslovakia 1,646 1,600 ? <10
Denmark 218 214 ? ?
France 11,740 12,000 250 2
Germany 5,707 5,700 100 2
Great Britain 11,560 4,800 *200 4
Greece 5,255 1,789 ? ?
Hungary 1,210 1,350 200 1S
Ireland 4,688 5,800 700 12
Italy 4,907 5,000 200 + 4 +
Netherlands - 1,100 10 1
Norway 102 - ? 10-20
Poland 4,040 4,040 ? C4
Portugal 531 761 ? ?
Romania 4,410 4,400 ? ?
Spain 10,210 6,645 1,452 22
Sweden 562 480 ? tg
Yugoslavia 6,347 6,400 ? ?
TOTAL 73,820 62,711 3,593 +
Notes: The FAO data (A) are for permanent pasture for 1988. The second column of figures (B) have been extracted from Dijk, G. van 1991. The
status of semi-natural grasslands in Europe. In: Goriup, P.D., Batten, L.A. and Norton, J.A. (Eds), The Conservation of Lowland Dry Grassland
Birds in Europe. The third column of figures (C) are for dry semi-natural grassland, all derived from van Dijk 1991, who used various sources.
The data are not all strictly comparable. This category of grassland omits wet or seasonally flooded grassland, upland grassland, acid grassland and
communities dominated by ericaceous species. Where separate figures were given for calcareous and neutral grassland, these have been combined.
* includes only lowland grassland in England and Wales.
Loss of semi-natural grassland has occurred because of the
enormous changes in agricultural methods. The use of
inorganic fertilizers and modern pesticides has vastly
increased productivity, at the expense of indigenous plants
and animals. The area under intensive cultivation in Europe
has increased dramatically since the 1950s. The trends have
been clearly illustrated for France by Lecomte and Voisin
(1991), from which Fig. 21.4 has been derived, showing
increase in the area of cereal crops and intensive agriculture
between 1970 and 1985.
In arid parts of Spain, Italy, the Balkans and Turkey, the
prevailing pattern of agriculture for most of the last century
or two has been to rotate grazing, arable cropping and
fallow. This has traditionally been done in small units of
land, creating a diverse landscape and normally one with a
good scatter of scrub and intermittently managed vegetation.
This pattern of land-use, which has been described by
Goriup (1988) as pseudosteppe, provides the ecological
conditions required by a very large number of plant and
animal species, many of which were originally part of the
steppe community. Birds, reptiles and small mammals are
generally mobile enough to accommodate this gently
shifting pattern of agriculture, so, even though it may be far
removed from natural or even semi-natural grassland,
pseudosteppe has become a very valuable wildlife
resource. The pseudosteppe style of land-use is now under
threat in Europe as people become increasingly dissatisfied
with this hard and relatively unrewarding way of life. The
trends are most apparent in the Mediterranean region but
similar pressures are having adverse effects on wildlife in
the western parts of the British Isles.
The Great Bustard Ofis tarda is an example of a bird
species formerly associated with extensive dry grasslands
286
but which was able to adapt and flourish in open
countryside maintained as a mosaic of grassland, arable and
fallow. Great Bustards will often display on grassland but
select arable land for their nest sites. Increasing dependence
on this kind of low intensity farming, places the birds at
great risk when agricultural intensification occurs,
particularly if there is increased application of pesticides
and fertilizers (Kollar, 1991). The Great Bustard has
undergone significant population decline through much of
its breeding range as a result. Other species particularly at
risk from this threat include the Little Bustard Tetrax tetrax,
Button Quail Turnix sylvatica, Sociable Plover Chettusia
gregaria and Demoiselle Crane Anthropoides virgo. In the
British Isles, the Corncrake Crex crex and the Chough
Pyrrhocorax graculus are very much at risk from similar
kinds of social and agricultural change (Goriup ef al.,
1991). The Large Blue butterfly Maculinea arion illustrates
similar trends for the invertebrates.
North America
The prairie zone of USA and Canada is inherently very
fertile and arable farming has been developed throughout
the region. A relatively small area managed to escape
ploughing and cropping through various accidents of history
and ownership. Much of the surviving prairie grassland is
therefore secondary in origin. The extent of ploughing and
agricultural improvement was greater in the western half of
the prairie zone, which is known as long-grass prairie, than
in the eastern short-grass prairie (Knopf, 1988). The rainfall
is higher in the former region, hence there is increased
agricultural production there.
A review of the survival of all the major vegetation types
in USA was carried out by Crumpacker et al. (1989). This
Grasslands
Table 21.5 Area of grassland habitats in USA (km?)
PNV TYPE* AREA OF AREA NAT % NATURAL
PNV IN USA OR S/N OR S/N IN USA
(1967) (1967)
Mesquite savanna 23,041 -
Mesquite-buffalo grass 70,406 51,236 72.9
Northern cordgrass prairie 3,779 2,562 67.8
Fayette prairie 7,696 4,302 55.9
Fescue oatgrass 3,565 2,110 59.2
Bluestem-Sacahuista prairie 41,457 9,908 23.9
Blackland prairie 48,461 15,216 31.4
Sea oats prairie 1,564 -
Grama-tobosa prairie 15,196 14,831 97.6
Southern cordgrass prairie 22,292 13,108 58.8
Palmetto prairie 11,273 8,251 73.2
Bluestem-Grama prairie 150,771 53,071 35.2
Wheatgrass-Grama-Buffalo grass 2,639 2,435 92.3
California steppe 51,973 15,903 30.6
Fescue-wheatgrass 20,985 5,561 26.5
Bluestem prairie 272,567 40,612 14.9
Aleutian meadows 12,730 -
Wheatgrass-bluegrass 36,845 25,238 68.5
Grama-buffalo grass 309,170 170,352 55.1
Wheatgrass-needlegrass shrubsteppe 27,258 25,867 94.9
Wheatgrass-needlegrass 253,707 161,414 63.8
Grama-needlegrass-wheatgrass 205,196 156,563 76.3
TOTAL 1,592,571 778,540
Source: Crumpacker, D.W., Hodge, S.W., Friedly, D. and Gregg, W.P. 1989. A preliminary assessment of the status of major terrestrial and
wetland ecosystems on Federal and Indian Lands in the United States. Conservation Biology 2(1):103-115.
Note: * PNV = Potential Natural Vegetation, NAT = natural, S/N = semi-natural.
is based on the concept of Potential Natural Vegetation
(PNV) which is defined as the vegetation that would, if man
were removed from the scene, exist in a region at the end
of the sequence of plant succession. Crumpacker et al. give
the area of each type considered to be still present in a
natural or semi-natural state in 1967 and calculate this as a
percentage of the potential natural vegetation. The
vegetation types which broadly equate to grassland and the
area of each said to have been present in 1967 are set out
in Table 21.5. The figures are based on small-scale maps,
however, and are subject to the usual problem of
amalgamating fragmented areas of grassland into a single
type. The actual areas of grasslands are therefore likely to
be inflated.
The total area of grassland PNV types would occupy about
17% of USA. This compares with a figure of 25.8% for the
proportion of USA occupied by permanent pasture
according to FAO data. The data in Crumpacker et al.
show that the grassland area which was considered to be
natural or semi-natural in 1967 occupied about 8% of USA.
As explained above, this is probably an inflated figure and
it is certainly likely that there has been further significant
loss since 1967. Table 21.5 shows that the ecosystems
which have been reduced by the greatest extent are the
bluestem prairie and the bluestem-Sacahuista prairie. A
considerable area of the once very extensive bluestem-
Grama prairie has also disappeared. The bluestem grasses
belong to the widespread genus Andropogon, which
constituted the dominant species over much of the
long-grass prairie. In some years, after fire they can grow
to a height of up to 2m, illustrating the potential
287
productivity of this habitat and hence the probable reason
for its disappearance.
Today, attempts are being made to recreate some of the
original prairie grasslands of the USA and Canada.
However, these newly-created prairie grasslands lack many
of the original indigenous species and the densities of plants
are thought to be much lower than they were in the natural
prairies. Nevertheless, the dominant grasses mirror the
original communities and the habitats can have a very
natural appearance. More information on the fate of the
prairies is given in the case studies on the Black-footed
Ferret and Canada.
Asia
The eastern part of the Asian steppes, particularly
Mongolia, still supports the most extensive and natural area
of the world’s great grasslands. Land-use practices have
been extraordinarily stable over the centuries, with low
intensity grazing being carried out by semi-nomadic
tribesmen. Agricultural practices have developed very little
and pesticides and fertilizers are largely irrelevant in this
shared system of grazing. The land is all in state-ownership
and people hold rights to drive their stock to wherever the
grazing is best. Over a vast area approaching the size of
western Europe, sparse grazing continues without fences to
confine stock. The only barriers are forests and mountain
ranges. Horses and sheep are the main grazers, with goats
and camels in the drier regions. Gazelles are still reasonably
-plentiful and are highly prized as a resource for hunting.
Changing social and political attitudes are now likely to
1. Biological Diversity
change traditional land-use practices.
The state of the grasslands is very much less satisfactory in
the former USSR than it is in Mongolia. A very high
proportion of the original steppe has been destroyed,
particularly in Kazakhstan and Uzbekistan. Very extensive
irrigation projects have been carried out, permitting
agricultural improvement on a huge scale. Before the
irrigation works were begun, large ploughs destroyed the
burrows of gerbils and other mammals. The main purpose
of this was to eliminate the gerbils before people had to
work on the land, because they carry a parasite which
causes leishmaniasis in man. Most of the irrigated land is
used for cotton crops, which are heavily treated with
insecticides and provide a very sterile environment for
wildlife.
The pressures on Indian grasslands have always been high
but they have been accelerating this century with the growth
in population (Majumdar and Brahmachari, 1988). Food
production has not increased in all areas, instead there has
been a major growth in production of cash crops. More
information on this is given in the case study on the Lesser
Florican.
Australia
In Australia the main problems in conserving grassland
habitats have arisen from alterations to traditional patterns
of burning. Aboriginal peoples had burned the vegetation in
a rotational system, timing the burns carefully in relation to
season and weather. This kept huge areas of the Australian
hinterland in a broadly open condition and increased
productivity for grazing animals. The fires were sufficiently
frequent to ensure that woody material did not accumulate
but well spaced enough to ensure that native plants and
animals could recover from the fire and take advantage of
the better growing conditions. Through the last hundred
years or so, as more of Australia has become settled, the
traditional burning patterns have been disrupted. Large
areas are burned annually, and very few of the native plants
and animals can cope with such a regime; pastures are
becoming increasingly dominated by introduced European
plants. Conversely, huge areas have been burnt very much
less frequently, and this has had two consequences: firstly,
species of grassland and other open habitats decline because
the vegetation becomes too thick, woody and tall, and,
secondly, when fires do happen they burn at a higher
temperature and are more destructive. In some national
parks and protected areas, efforts are now being made to
return to the traditional burning patterns (Boekel, 1990),
with the direct help and involvement of the indigenous
people.
CASE STUDIES
The following case studies illustrate the types of threat
facing declining species of grassland-adapted fauna, and
those affecting the habitat as a whole.
The Lesser Florican as an indicator of grassland loss in
India
The Lesser Florican Sypheotides indica is the smallest and
288
formerly most widespread of the three bustard species
endemic to the Indian subcontinent. The majority of the
birds both breed and winter in dry grassland, though there
is some migration to areas where rainfall has increased food
availability. Until the 1980s, conservationists had been
concentrating most of their concern on the Great Indian
Bustard Ardeotis nigriceps. The Lesser Florican was
thought to be common and widespread. It was a popular
bird for the table and large numbers were shot by
sportsmen and caught in nets. Indeed, until 1980 the species
was still officially sanctioned as legal prey for hunters.
Field surveys were made in four separate areas of India
which had been reported as strongholds of Lesser Floricans
within recent decades (Goriup and Karpowicz, 1985). One
of these, the Tungabadhra Wildlife Sanctuary, near Bellary
in Karnataka, southern India, no longer supported any
grassland as a large reservoir had been constructed and all
the adjacent land in the valley had been developed for
agriculture. In the Jaipur area of northern India, grassland
was still present in some quantity but was overgrazed and
generally unsuitable for Lesser Florican. The birds there
were reported to have become very much rarer in the last
20 years. The Deccan plateau of central India had been
regarded as the core of the Lesser Florican’s range but the
great majority of the semi-natural grassland had been
converted to rice paddies. When the paddies are prepared
for sowing the only remaining grass is on the embankments
(nallas), and the villagers use these for trapping
game-birds, including Lesser Floricans. Not surprisingly,
the birds have become very much less frequent and the
prospects for their survival in this area are very poor
(Goriup and Karpowicz, 1985).
Goriup and Karpowicz concentrated the majority of their
fieldwork in the Jamnagar district of the Kathiawar
Peninsula in north-west India. Here the remnants of the
once extensive grassland are under the control of the
Gujerat Forest Department, and they occur as discrete
patches known as vidis. The policy for these areas is that
they should be cut for hay and kept free from grazing stock
throughout the year. It was found, however, that the habitat
had deteriorated in many of the vidis, with scrub invasion
in some and cattle or buffalo grazing in others. Out of 50
vidis visited, seven were found to support Lesser Floricans,
and there was a total of only 22 individuals within them. A
more extensive survey of suitable areas in the region failed
to produce any more birds. It was apparent that Lesser
Floricans have not managed to adapt to new habitats such
as fields of groundnuts. Historical data on the land-use of
the Jamnagar district showed that, although the area
producing food crops had fallen from 34% to 18% between
1906 and 1981, the total cultivated area had risen from 45%
to 70% because of the increase in the district of cash crops,
principally groundnuts, cotton and sugar cane. The
agricultural pressures on the region are clearly very intense
and the prospects for Lesser Floricans and other grassland
species are not encouraging.
Grasslands in Poland
The extent and condition of grasslands in Poland have been
effectively reviewed by IUCN (1991), much of the
information having been drawn from Denisiuk (1990).
Poland occupies 312,000km7, of which 13% (or 40,400km”)
is grassland. This area has been reduced markedly in recent
years. The great majority of the semi-natural grassland is in
the major river valleys and would be classified as either
damp or wet. True steppe grasslands are now very rare in
Poland and are confined to steep south-facing slopes. Dry
species-rich, semi-natural grassland has become rare
because the level of fertilizer application in the country as
a whole is very high and well above the threshold for
maintaining floristic diversity. Quite a high proportion of
the dry grasslands is in fact former peat bogs which have
been drained and grazed. Some of these drained sites have
been brought under cultivation but the current estimate is
that 82% of their area is now maintained as meadows, and
these are classed as dry grassland. Management of this land
can be particularly difficult because as the peat dries it
oxidises and shrinks, thereby lowering the land surface,
which then requires draining with new deep ditches. Other
areas become too dry, with the result that yields fall and the
grassland becomes uneconomic. This is one reason why
there are about 10,000km? of abandoned farmland in
Poland.
An important characteristic of rural Poland is that it is held
in the form of very small farms, averaging only Sha. Large
farms were not formed in the same way as in other eastern
European countries because of resistance to collectivisation
after the Second World War. This results in a large area of
little-used boundary land, which is often of value for
wildlife, sometimes including grassland species. This land
and the abandoned farmland is, however, much more likely
to benefit the more adaptable species which require cover
and woodland edge.
The pressures on grassland flora and fauna have been
considerable in recent years. Agricultural intensification,
particularly increasing fertilizer use, is affecting all parts of
the country and more of the dry grassland has been brought
under cultivation. The area of wet grassland has been
reduced by drainage from 36% to 23% of the area of all
grasslands between 1973 and 1988.
The economic pressures on Poland’s farmers are forcing
them either to improve their agricultural methods, through
such measures as drainage, fertilizer use, irrigation and
switching from hay to silage, or to abandon the land
altogether. Further areas of grasslands have been lost to
afforestation, and it seems likely that this trend will
increase. For the foreseeable future, the best way of
maintaining the flora and fauna of Poland’s grasslands
would appear to be through the establishment of national
parks and nature reserves, rather than through changing the
direction of agricultural development.
The Meadow Viper
The Meadow Viper Vipera rakosiensis is the smallest and
least venomous of the European vipers. It is also the rarest,
having long been restricted to a specific lowland grassland
habitat in central Europe. It is found in both wet and dry
grasslands but is particularly associated with the interface
between the two. Sites providing large tussocks and ant-hills
are favoured, especially if there is varied topography
offering a range of soil moisture conditions and
289
Grasslands
temperatures. The past distribution of the Meadow Viper is
not well known but it is clear that the subspecies has
undergone a very severe decline. The situation in each
country is as follows:
Hungary. Still present in the Great Plain pusztas between
the rivers Danube and Titza, south of Budapest. Elsewhere
the habitat is severely fragmented anc under pressure from
agricultural improvement. Several sites have been lost as a
result of grazing and grubbing by geese and pigs. One 12ha
meadow has been protected in the Little Plain Hansag.
Romania. Recently became extinct following ploughing
and agricultural improvement of the Stipa (feather grass)
meadows with which it had long been associated.
Austria. It was formerly common in the sandy basins of
Vienna and Neusiedler but agricultural improvement has
destroyed almost all of its habitat. About 17ha of meadow
are now protected and suitably managed but the Meadow
Viper is generally thought to be extinct in Austria.
The causes of the decline of the Meadow Viper are very
clear:
Killing for bounty (when the species was more
common), especially in Austria
Land drainage and subsequent use for vineyards
Arable farming
Forestry
Application of fertilizer and pesticides
Increase in the frequency of mowing
Rearing of pheasants, which are predators of young
snakes
Collecting, for museums and private collections.
The decline of the Meadow Viper provides an illustration of
the effects of the pressures on central European meadows
which were formerly managed in a casual or inefficient
way. It is an example of a species which has been unable to
adapt to the reduction of structural complexity and
biodiversity in its grassland habitat.
The Canadian prairies
Natural grasslands were concentrated in the southern parts
of the three prairie provinces, Alberta, Saskatchewan and
Manitoba. Their characteristics and development have been
reviewed by Mondor and Kun (1982). The Canadian
prairies occupied the northern part of the north temperate
zone and are situated to the south of a large zone of aspen
parkland. The area of woodland was increased by farmers
who planted trees for shelter but many of these farms have
since been abandoned, thus allowing woodland to spread.
There are still huge open plains, however, supporting the
typical long-grass and short-grass prairie habitats.
At the time of settlement in the 19th century the extent of
open grassland in the prairie zone of Canada was probably
in the range of 360,000-400,000knr. This had been reduced
to about 80,000km? by 1982 and was reported to be
undergoing conversion to arable at a rate of approximately
500km? per year (Mondor and Kun, 1982). The FAO figure
for permanent pasture in the whole of Canada in 1985 was
1. Biological Diversity
325,000km?, so obviously a high proportion of this is
secondary and most of it will be sown or agriculturally
improved pasture of little biological interest.
Cattle ranching began in the Canadian prairies in the 1870s
and increased so rapidly that there was acute shortage of
land by the 1890s. Mismanagement, overgrazing and hard
winters forced most stockmen out of business by the early
years of the 20th century and cereal production became the
dominant land-use. Today open-range cattle ranching
survives only in south-eastern Alberta and in an adjacent
area in Saskatchewan. Elsewhere cattle grazing takes place
as part of an arable rotation system and recent decades have
seen very large increases in stock numbers.
The millions of bison which roamed the plains were
reduced to a low ebb of approximately 1,100 by 1889.
Other mammals were also reduced to very low levels,
primarily through hunting for food, notably the Elk Cervus
elaphus and Pronghorn Antelope Antilocarpa americana.
The latter was estimated to have numbered 50 million
animals, mostly on the open prairies, but by 1915 it had
been reduced to only a few herds in south-eastern Alberta
and south-western Saskatchewan. On the other hand,
increased grazing and shorter grass benefited small
mammals such as Pocket Gophers Geomys spp.,
Richardson’s Ground Squirrel Spermophilus richardsoni and
the Black-tailed Prairie Dog Cynomys ludovicianus. This
resulted in a dramatic increase in numbers of coyotes Canis
latrans (their predators), which became a major pest.
Poisoning and trapping were undertaken on a large scale,
and the Black-tailed Prairie Dog is now confined to only a
few small colonies in south-western Saskatchewan.
The losses of wildlife were on such a scale that
conservation efforts began at an early date. Legislation to
control hunting was passed between 1905 and 1915, and the
Buffalo National Park was established in 1908, a 440km?
refuge of long-grass prairie where a herd of bison still
survived. By 1922 this herd had increased to over 6,000
individuals, more than the park could support. Despite
vigorous efforts to find other areas to receive surplus
animals, none were found and 2,000 buffaloes were
slaughtered in 1923. Buffalo National Park did not prove
successful for the conservation of Pronghorn Antelope and
other grassland sanctuaries were established as national
parks primarily for this purpose, namely Nemiskam
(21km?), Wawaskesy (154km?) and Menissawok (44km?).
These measures achieved their objective and allowed the
Pronghorn to multiply to such an extent that it was no
longer threatened in any way. Surprisingly, the three
national parks were considered redundant and they were
decommissioned between 1930 and 1947. Since then there
have been extraordinarily protracted negotiations between
the Federal and Provincial governments to establish other
protected areas. A formal agreement to establish a
grasslands national park was signed between Saskatchewan
and the Federal government in 1981. In 1988,
Saskatchewan transferred to the Federal government all the
Tights over a core area of 187km?, but the balance of the
proposed park, an area of 719km?, may not be transferred
until the year 2021. If all the intentions are honoured this
will form a magnificent example of the prairie habitat, but
290
prospects for survival of semi-natural grasslands outside this
area are very poor.
The Black-footed Ferret - a species whose decline was
not linked to habitat loss
The Black-footed Ferret Mustela nigripes formerly occupied
a very large area within the central prairie zone of North
America, from Alberta to Arizona. It was primarily
associated with colonies of prairie dogs Cynomys spp. but
was occasionally found in the burrows of ground squirrels.
Ferrets were hunted by native Americans for their pelts but
the main reason for the dramatic population decline in the
last 100 years or so is considered to have been poisoning of
prairie dog colonies (Schreiber et al., 1989). This resulted
in very high mortality of ferrets, presumably as a direct
effect of poisoning and through loss of their main prey. It
appears, however, that diseases such as canine distemper
have also had a very significant impact on the reduced
populations. In 1920, numbers were estimated at over
500,000 (Clark, 1987) but the catastrophic decline
continued. By 1937 the species was extinct in Canada and
by 1950 it was feared to be so in USA as well. This was
despite the fact that there was no shortage of suitable habitat
- as much as 400,000km?”, it was thought in 1970.
In 1964 a Black-footed Ferret population was discovered in
prairie dog colonies in South Dakota but by 1974 this
population had disappeared. In 1981 another population was
found near Meeteetsee, in Wyoming, confined to a total
area of about 30km?, but scattered over 130km*. Numbers
had probably fluctuated around 100 or so individuals for
about 50 years. There was a peak of 129 animals in 1984
but canine distemper reduced this to only 12 in 1985. It was
considered that captive breeding provided the only hope for
the species, so 24 individuals were taken into captivity
between 1985 and 1987. The wild population did not
survive. Initially disease caused further losses in captivity
but this problem has been overcome by strict isolation and
quarantine. Captive breeding has now raised numbers
substantially. Plans for reintroduction to suitable areas are
being drawn up. A number of large prairie sites are being
managed appropriately and, outside these, farmers are being
paid to protect their prairie dog colonies. The Black-footed
Ferret is clearly a highly specialised grassland species
unable to adapt to new conditions or to switch to other
prey. It stands little chance of survival without a fully
researched and properly resourced conservation strategy.
The Steppe Marmot - adapting to the changing
conditions
The Steppe Marmot Marmota bobac was found in the short
grassy steppes in eastern Europe, from Hungary to the
Urals. The former very extensive range of this burrowing,
hibernating herbivore was steadily reduced by cultivation
and hunting until it reached a low point in the 1940s and
1950s (Bibikov, 1991). Only a few thousand survived, in a
huge area between the Ukraine and the Urals. They were
restricted to a handful of rather unsuitable pastures in
valleys or areas where dissected relief and other factors
prevented ploughing. In recent decades the Steppe Marmot
has made a spectacular recovery, partly through
reintroduction, protection and conservation measures but
also as a result of adaptation by the species. Following rural
depopulation, the marmots began to colonise abandoned
farmsteads and villages and are even found in unused parts
of occupied villages. There they find a variety of food
plants and good conditions for burrowing. The population
in USSR was thought in 1991 to be around 250,000
individuals, representing at least a ten-fold increase since
the 1940s. A high proportion of these are now using
farmland, including cultivated ground, and they have clearly
adapted to using a wider variety of food items. Until the
middle of this century the marmot would have been
regarded as one of the species most typical of steppe
grasslands. However, when pressure on the habitat became
acute, the marmot began to demonstrate an inherent
capacity for adaptation and this has resulted in a
transformation of its prospects.
Native grassland and grassy woodlands in Victoria,
Australia
Today there is little native grassland in the state of Victoria
but before European settlement no less than 34% of the area
had supported either grassland or grassy woodland. The
topic has been reviewed in a convincing Conservation
Strategy by Baker-Gabb and Lunt (1990).
Baker-Gabb and Lunt do not distinguish between grassland
and grassy woodland, but produce abundant evidence that
together they constitute the most threatened ecosystem in
Victoria. For instance, at least 125 of Victoria’s 866 rare
and threatened plant species occur in these habitats, they
include 28 composites, 14 legumes and nine orchids. No
less than 31% of the endangered plant species are confined
to these habitats, mostly in only one area of the state, while
eight plant species that used to occur are now extinct in the
state. An even greater number (26) of vertebrate species
have become extinct. Indeed, of the 152 species of extinct,
endangered/vulnerableand threatened species of vertebrates
in Victoria, no fewer than 61 (40%) are associated with
grasslands and grassy woodlands.
The rich soils of the native grasslands of Victoria were very
attractive to early settlers anxious to make a living from
agriculture. Ploughing, re-seeding and overstocking with
sheep and cattle had disastrous effects on the native fauna
and flora. The habitat loss was too complete for significant
areas to be incorporated into the major national parks which
have been established since the early 1970s. Today the
majority of conservation areas which do contain grassland
are small, isolated and surrounded by agricultural or urban
land. Thus the network of national parks has done relatively
little to conserve grasslands, since only 0.3% of the original
area has received protection and little of the original
diversity is represented.
References
Atlay, G.L., Ketner, P. and Duvigneaud, P. 1979. Terrestrial primary
production. In: Bolin, B. (Ed.), The Global Carbon Cycle. Wiley,
Chichester.
Baker-Gabb, D.J. and Lunt, I.D. 1990. Conservation Program for
Native Grasslands and Grassy Woodlands in Victoria. Department
of Conservation and Environment. Melbourne.
291
Grasslands
Bibikov, D. 1991. The steppe marmot - its past and future. Oryx
25:45-49.
Boekel, C. 1990. Traditional aboriginal land management practices in
Australian national parks. Parks 1(1): 11-15.
Cabrera, A. 1970. Flora de la Provincia de Buenos Aires: gramineas.
Instituto Nacional de Tecnologia Agropecuara, Buenos Aires.
624pp.
Clark, T.W. 1987. Black-footed ferret recovery: a progress report.
Conservation Biology 1(1):8-10.
Clements, F.E. and Shelford, V.E. 1939. Bioecology. Wiley, New
York.
Crumpacker, D.W., Hodge, S.W., Friedly, D. and Gregg, W-P. 1989.
A preliminary assessment of the status of major terrestrial and
wetland ecosystems on Federal and Indian Lands in the United
States. Conservation Biology 2(1):103-115.
Denisiuk, Z. 1990. Lowland grasslands in Poland - their natural
resources, managementand protection. In: The Lowland Grasslands
of Eastern Europe. YUCN, Gland.
Dijk, G. van 1991. The status of semi-natural grasslands in Europe.
In: Goriup, P.D., Batten, L.A. and Norton, J.A. (Eds), The
Conservation of Lowland Dry Grassland Birds in Europe,
Proceedings of an International Seminar held at the University of
Reading 20-22 March 1991.
Duffey, E., Morris, M.G., Sheail, J. and Wells, T.C.E. 1974.
Grassland Ecology and Wildlife Management. Chapman and Hall,
London.
Folster, H. and Huber, O. 1984. Interrelaciones Suelos-vegetacién en
el Area de Galipero, Territorio Federal Amazonas, Venezuela
(Series Informes Tecnicos DGSIIA/IT/144), Ministerio del
Ambiente y de los Recursos Naturales Renovables.
Food and Agriculture Organization 1987. World Crop and Livestock
Statistics 1948-1995; Area, Yield and Production of Crops;
Production of Livestock Products. FAO, Rome.
Goriup, P.D. 1988. The avifauna and conservation of steppic habitats
in western Europe, North Africa and the Middle East. In: Goriup,
P.D. (Ed.), Ecology and Conservation of Grassland Birds. \CBP
Technical Publication No.7. ICBP, Cambridge.
Goriup, P.D., Batten, L. and Norton, J. (Eds) 1991. The Conservation
of Lowland Dry Grassland Birds in Europe. Proceedings of an
International Seminar held at the University of Reading 20-22
March 1991.
Goriup, P.D. and Karpowicz, Z. 1985. A review of the past and recent
status of the lesser florican. Bustard Studies 3:163-182.
Goriup, P.D. and Schulz, H. 1991. Conservation Management of the
White Stork - an international need and opportunity. ICBP
Technical Publication No.12. ICBP, Cambridge.
Grimmett, R.F.A. and Jones, T.A. 1989. Important Bird Areas in
Europe. ICBP Technical Publication No.9. ICBP, Cambridge.
Groves, R.H. 1981. Australian Vegetation. Cambridge University
Press, Cambridge.
Huber, O. 1987. Neotropical savannas: their flora and vegetation. Tree
2(3).
TUCN 1991. The Lowland Grasslands of Central and Eastern Europe.
Cambridge.
Kollar, H.P. 1991. Status of lowland dry grasslands and great bustards
in Austria. In: Goriup, P.D., Batten, L. and Norton, J. (Eds), The
Conservation of Lowland Dry Grassland Birds in Europe.
Proceedings of an International Seminar held at the University of
Reading 20-22 March 1991.
Knopf, F.L. 1988. Conservation of steppe birds in North America. In:
Goriup, P.D. (Ed.), Ecology and Conservation of Grassland Birds.
ICBP Technical Publication No.7. ICBP, Cambridge.
Knystautas, A. 1987. The Natural History of the USSR. Century,
London.
Lecomte, P. and Voisin, S. 1991. Dry grassland birds in France:
status, distribution and conservation measures. In: Goriup, P.D.,
Batten, L. and Norton, J. (Eds), The Conservation of Lowland Dry
Grassland Birds in Europe. Proceedings of an International Seminar
held at the University of Reading 20-22 March 1991.
Lee, J. 1990. Land use trends and factors influencing change in future
land use in EC-12. European Agrarian Youth Congress, Groningen.
1. Biological Diversity
Majumdar, N. and Brahmachari, G.K. 1988. Major grassland types
and their bird communities: a conservation perspective. In: Goriup,
P.D. (Ed.), Ecology and Conservation of Grassland Birds. ICBP
Technical Publication No.7. ICBP, Cambridge.
Menaut, J-C, 1983. The vegetation of African savannas. In: Bouliére,
F. (Ed.), Tropical Savannas. Ecosystems of the World, 13.
Elsevier, Amsterdam.
Moore, R.M. 1970. Australian Grasslands. Alexander Bros.,
Melbourne.
Mondor, C. and Kun, S. 1982. The long struggle to protect Canada’s
vanishing prairie. Ambio 2:286-291.
Olson, J.S., Watts, J.A. and Allison, L.J. 1983. Carbon in Live
Vegetation of Major World Ecosystems. Oak Ridge National
Laboratory, for US Department of Energy, Washington.
Polunin, O. and Walters, M. 1985. A Guide to the Vegetation of
Britain and Europe. Oxford University Press, Oxford. 238pp.
292
Rogers, D.L. and Randolph, S.E. 1988. Tsetse flies in Africa, bane or
boon? Conservation Biology 2(1):57-65.
Sarmiento, G. 1983. In: Bouliére, F. (Ed.), Tropical Savannas.
Ecosystems of the World, 13. Elsevier, Amsterdam. Pp.245-288.
Schreiber, A., Wirth, R., Riffel, M. and Van Rompaey, H. 1989.
Weasels, Civets, Mongooses and their Relatives - An Action Plan
for the Conservation of Mustelids and Viverrids. IUCN, Gland.
Stuart, S.N. and Adams, R.J. 1990. Biodiversity in Sub-Saharan
Africa and its Islands. Occasional Papers of the IUCN Species
Survival Commission No.6. Oxford.
Whittaker, R.H. and Likens, G.E. 1975. The biosphere and man. In:
Leith, H. and Whittaker, R.H. (Eds), Primary Productivity of the
Biosphere. Springer-Verlag, Berlin.
Abridged from a consultancy report written by Richard J.
Hornby of the Nature Conservation Bureau (UK).
22. WETLANDS
The term ‘wetlands’ groups together a wide range of inland,
coastal and marine habitats which share a number of
common features. The Ramsar Convention defines wetlands
as "areas of marsh, fen, peatland or water whether natural
or artificial, permanent or temporary, with water that is
static or flowing, fresh, brackish or salt, including areas of
marine water the depth of which at low tide does not exceed
six metres".
However, in spite of the apparent clarity of this definition,
the classification of wetlands is fraught with problems.
There are an enormous variety of wetland types, even the
broadest grouping of habitat types according to their basic
biological and physical characteristics gives 30 categories of
natural wetlands and nine man-made ones (Dugan, 1990).
In addition, wetlands are highly dynamic, changing with the
seasons and over longer periods of time and it is frequently
difficult to define their boundaries with precision. As a
result, estimates of area of wetland vary considerably and
it is not always clear what particular kinds of habitat are
being discussed.
GLOBAL EXTENT AND DISTRIBUTION OF
WETLANDS
Two recent papers (Aselmann and Crutzen, 1989; Matthews
and Fung, 1987) give estimates of the global distribution of
wetlands, but both are concerned primarily with methane
production and they do not, therefore, include any salty
areas as these do not emit methane to any great extent.
Although very different methods were used to calculate the
area of natural freshwater wetlands, the two estimates are
very similar. Matthews and Fung (1987) combined three
independent data sources: the first was a global vegetation
database classified with the UNESCO system; the second
was a global database digitised from FAO soil maps and the
third was a global inundation data base compiled from
Operational Navigation Charts. They concluded that
5.3 million km? of wetland remain. Aselmann and Crutzen
(1989), using information from Gore (1983) and a variety
of map sources, estimated that 5.7 million km? of
Wetlands
freshwater wetlands existed. A comparison of the
distribution of these wetlands according to the two reports
is given in Table 22.1.
Both authors also show the distribution of the different
types of wetland along 10° latitude belts, but the categories
used in the two papers are very different. Fig. 22.1 shows
this distribution of types according to Aselmann and
Crutzen (1989) and Table 22.2 indicates their estimates of
wetlands in various countries or regions. Their definitions
of each vegetation type are given below.
Bogs
Peat-producing wetlands in moist climates where organic
matter has accumulated over long periods. Water and
nutrient input is entirely through precipitation. They are
acid and nutrient deficient. Sphagnum moss typically
dominates the vegetation.
Fens
Peat-producing wetlands which are influenced by soil
nutrients flowing through the system. Grasses and sedges,
with mosses, are the dominant vegetation. These are
generally more prolific than bogs.
Swamps
Forested freshwater wetlands on waterlogged or inundated
soils where little or no peat accumulation occurs.
Marshes
Herbaceous mires with vegetation commonly dominated by
grasses, sedges or reeds. They may be either permanent or
seasonal. Salt marshes have been excluded.
Floodplains
Periodically flooded areas along rivers or lakes. They show
considerable variation in vegetation cover.
Shallow lakes
Open water bodies a few metres in depth. Regional extent
and distribution of wetlands
Table 22.1 Comparison of two estimates of global wetland area* along 10° latitude
belts
LATITUDES
NORTH SOUTH
SOURCE 80-70 70-60 60-50 50-40 40-30 30-20 20-10 10-0 0-10 10-20 20-30 30-40 40-50
(A) 122 1355 1235 319 128 94 276 431 484 360 333 132 3
(B) 130 1481 1445 276 156 49 85 488 1062 393 85 29 10
Difference 8 126 210 -43 28 -45 -191 57 578 33 -248 -103 7
(A) - (B)
Source: (A) = Matthews, E. and Fung, I. 1987. Methane emission from natural wetlands: global distribution, area and environmental characteristics
of sources. Global Biogeochemical Cycles 1(1):61-68.; (B) = Aselmann, I. and Crutzen, P.J. 1989. Global distribution of natural freshwater
wetlands and rice paddies, their net primary productivity, seasonality and possible methane emissions. Journal of Atmospheric Chemistry 8:307-358.
Note: * In 1,000km?.
293
1. Biological Diversity
Latitudinal distribution of natural wetlands
Figure 22.1
sousieW 9 sud ‘sbog iz sae] MO|eEUS oC surejdpoo|4 | sousieyy LL sdwems ey sud
vb ct
(2wuy SUONIW) SpUBNaM [eINJeN JO Bary
L 80 90 v0
294
Wetlands
Table 22.2 Global freshwater wetland areas*
REGION BOGS FENS SWAMPS
‘USSR’ 917 531 25
Europe 54 93 1
Near East - - =
Far East - = 11
China 11 - 3
Southeast Asia 197 - 44
Aust/NZ 2 3 1
Africa - - 85
Alaska ? 250 - 400 ?
Canada 673 531 14
USA** 13 = 80
C America - - 15
S America - = 851
TOTAL 1,867 1,483 1,130
MARSHES FLOODPLAINS LAKES TOTAL
39 = 1,512
4 1 1 154
8 = 8
= 11
18 32
- = 241
- 9 = 15
ey/ 174 39 355
? ? ? (325)
44 = 1,268
40 95 228
2 1 = 18
62 543 68 1,524
274 823 114 5,691
Source: Aselmann, I. and Crutzen, P.J. 1989. Global distribution of natural freshwater wetlands and rice paddies, their net primary productivity,
seasonality and possible methane emissions. Journal of Atmospheric Chemistry 8:307-358.
Notes: * In 1,000km; ** excluding Alaska.
REGIONAL EXTENT AND DISTRIBUTION
WETLANDS
OF
A recent survey of the world’s wetlands (Finlayson and
Moser, 1991) includes maps of the distribution of selected
major wetlands; some of these maps have been incorporated
into Fig. 22.2. The important wetlands of Asia are shown
in Fig. 22.3 (data from Scott and Poole, 1989). In many
cases where regional estimates of wetlands areas have been
calculated, they are very different from those given by
Aselmann and Crutzen (1989). This is because of the
inclusion of salty areas and, no doubt, because different
definitions of wetlands and different methods of estimation
were used in each set of calculations.
Europe and the Mediterranean Basin
Europe and the Mediterranean are so densely populated and
have had such long histories of civilization and
industrialisation that there are only a few entirely natural
wetlands left in this area. Human interference has been less
severe in parts of Iceland and the northern European taiga
and tundra, but in most other regions the wetlands have
either gone or are threatened. For instance, by the end of
the 1970s, 10% of France’s wetter areas and 60% of those
of the UK and the Netherlands had been drained (Finlayson
and Moser, 1991). There has, however, been extensive
creation of artificial wetlands such as reservoirs, fishponds
and gravel pits. In Tunisia, for example, 224km? of open
water have been created while, since 1881, 190km? of
natural wetlands have been lost (Finlayson and Moser,
1991). However, most countries have now joined the
Ramsar Convention and the rate of destruction of the
wetlands may at least be slowing down. Data for all the
European and Mediterranean Ramsar sites reveal that only
58 of the 318 wetlands are definitely not threatened in some
way.
North America
Canada is estimated to hold 24 % of all the world’s
wetlands, occupying over 1.27 million km? (Finlayson and
Moser, 1991). The original wetland area of the
295
conterminous USA (excluding Alaska and Hawaii) may
have been around 890,000km’, of which only 47 % or
thereabouts remain (Dahl, 1990). There were a further
690,000km? in Alaska and Hawaii, with only a very small
percentage in the latter state. Overall, it is estimated that
around 1.11 million km? of wetlands remain in the whole of
the USA (Dahl, 1990). Fig. 22.5 shows the distribution of
wetland in the USA about 200 years ago and in the 1980s.
The percentage loss of wetland in each state is also shown
in Fig. 22.5.
Latin America and the Caribbean
Many of the wetlands in South America are in an almost
pristine state. In contrast, most of the habitats, including the
wetlands, in the Caribbean have been intensively exploited.
The state of the natural habitat in Central America and
Mexico is intermediate, with a fairly large area of wetlands
remaining.
The wetlands of South America can be subdivided into three
major systems: those of the Pacific lowlands, those of the
Andean chain and those of the Atlantic-Caribbean lowlands
to the north and east of the Andes. The Chilean Fjordland
in the Pacific lowlands includes around 55,000km? of
wetlands (Finlayson and Moser, 1991). The largest wetland
in the Andes region is the freshwater lake, Lake Tota, in
Colombia. Also in this area is the Chilean Lake District
covering some 3,000km? of wetland. In the lowlands of the
Atlantic-Caribbean region, the delta of the River Orinoco
covers an area of around 30,000km?, while that of the
Amazon River covers about 35,000km?. Also in this area is
the Pantanal, covering some 200,000km”, which is one of
the largest floodplains in the world.
Many of the Caribbean islands have important wetlands,
mostly coastal lagoons, mangrove swamps and inter-tidal
mudflats, but there are also some freshwater lakes in old
volcanic craters. Many of the flora and fauna found on the
islands are endemic.
The Usumacinta Delta is the most extensive wetland on the
Gulf coast of Mexico, covering around 10,000km7.
1. Biological Diversity
Figure 22.2 General distribution of world wetland areas
2000 4000 6000
jo)
296
Wetlands
Table 22.3 Wetlands described in A Directory of Asian Wetlands
COUNTRY NUMBER OF SITES AREA OF SITES (km?)
Bangladesh 12 67,700
Bhutan 5 85
Brunei 3 1,380
China 192 163,000
Hong Kong 3 119
India 93 54,700
Indonesia 137 87,800
Japan 85 4,750
Cambodia 4 36,500
Korea, Dem People’s Rep 15 3,220
Korea, Rep 21 1,070
Laos 4 2,220
Malaysia 37 31,200
Mongolia 30 15,500
Myanmar 18 54,900
Nepal ilz/ 356
Pakistan 48 8,580
Papua New Guinea 33 101,000
Philippines 63 14,100
Singapore 7 2
Sri Lanka 41 2,740
Taiwan 12 84
Thailand 42 25,100
Viet Nam 25 58,100
TOTAL 947 734,200
Source: Scott, D.A. and Poole, C.M. 1989. A Status Overview of Asian Wetlands. No.53. AWB, Kuala Lumpur, Malaysia.
Africa
Wetlands cover one per cent of Africa’s total surface area
(at least 345,000km7?). In Equatorial Africa, the three largest
wetland systems are: the Zaire swamps (covering
80,000km7), the Sudd in the Upper Nile (over 50,000kn?r)
and the wetlands of the Lake Victoria Basin (about
50,000km’). The floodplains of the Niger and Zambezi
Rivers, the Chad Basin (around 20,000km?) and the
Okavango Delta (16,000km?) are also major wetland areas.
There are also a further 12,000km? of wetland in southern
Africa.
Asia and the Middle East
It has been estimated that there are some 830,000km? of
peat bogs and swamps in the USSR and about 900,000km?
of marshy ground subject to seasonal flooding (Finlayson
and Moser, 1991). In the Middle East, the most extensive
wetlands occur in Iraq, where the Tigris and Euphrates
Rivers create a vast complex of shallow lakes and marshes
covering about 15,000km?. It is estimated that there are
around 1.2 million km? of wetlands, excluding permanent
rice paddies, in the region covered by the Directory of
Asian Wetlands (Scott and Poole, 1989). This Directory
gives information on 947 of the most important wetlands,
covering 734,000km?, and their distribution by country is
shown in Table 22.3.
Australasia and Oceania
The major wetland types in this region are seagrass
meadows, mangrove swamps, coastal salt marshes and flats,
monsoonal freshwater floodplains, southern and inland
swamps, lakes, river and creek channels and bogs
297
(Finlayson and Moser, 1991).
The seagrass meadows off the coast of Australia are some
of the largest in the world. Mangroves are another
important wetland habitat in the region. They cover
12,000km? in Australia, 9,250km* in Papua New Guinea
and about 28,500km? in Irian Jaya, but only 640km? in the
Oceanic islands and a small area in New Zealand (Finlayson
and Moser, 1991). Salt marshes occupy about 9,200km* in
Australia and are also found in New Zealand. Extensive
herb, woodland and forested freshwater floodplains occur
in Australia and there are also numerous swamps along the
rivers, both here and in New Zealand. For instance, at the
confluence of the Lachlan and Murrumbidgee Rivers there
are nearly 1,500km?* of wetlands and another 400km* of
swamp along the Macquarie River in New South Wales
(Finlayson and Moser, 1991). Lakes, both saline and
freshwater, permanent and temporary, are found throughout
the region.
VALUES AND THREATS
Wetlands serve a wide variety of functions, including flood
control, water purification, shoreline stabilisation and the
control of erosion. They also support vast numbers of fish
and other wildlife and numerous people depend on them for
their livelihood. Table 22.4 lists these values and indicates
which types of wetland provide them. This topic is
developed further in Part 2.
The services provided by wetlands have tended to be taken
for granted and, as a result, maintenance of natural
wetlands has received low priority. Indeed, in many cases
the drainage of wetlands has been seen as an advantage,
with the benefits far outweighing the costs, whereas, in
1. Biological Diversity
fact, the opposite often turns out to be nearer the truth.
Table 22.5 lists the general causes of wetland loss, in
particular habitat types. Major threats to wetlands in the
Neotropics are summarised in Table 22.6.
A listing of general threats to wetlands in Asia is given in
Table 22.7, with a more detailed country-specific analysis
in Table 22.8.
In Asia, there are regional differences in the frequency of
occurrence of particular types of threat. In East Asia,
human settlement and encroachment occur in 20% of the
212 sites on which data have been gathered, while
reclamation for urban and industrial development and
pollution are both reported in 18% of the sites. In Southeast
Asia, the most common threats are hunting and the
disturbance associated with it (occurs at 42% of the 331
sites for which information is known), disturbance from
human settlement and encroachment (34%), commercial
logging and forestry (30%), wood-cutting for domestic use
and drainage for agriculture (both 27%). In South Asia,
hunting and its associated disturbance is, again, the most
common threat, occurring at 39% of the 191 sites; fishing
Table 22.4 Wetland values
Functions
Groundwater recharge
Groundwater discharge
Flood control
Shoreline stabilisation/Erosion control
Sediment/toxicant retention
Nutrient retention
Biomass export
Storm protection/windbreak
Micro-climate stabilisation
Water transport
Recreation/Tourism
Products
Forest resources
Wildlife resources
Fisheries
Forage resources
Agricultural resources
Water supply
Attributes
Biological diversity
Uniqueness to culture/heritage
Estuaries (without
is also a threat in many areas (32%). The other most
common threats in this region are overgrazing by domestic
livestock (27%), pollution (26%) and degradation of
watersheds with soil erosion and increased siltation (25%).
The most seriously threatened wetlands in Asia have been
shown in Fig. 22.4; those marked in black are considered
to be already too degraded to merit any special conservation
effort (Scott and Poole, 1989). They are also listed in Table
22.9, where the degraded sites are marked with an asterisk.
LOSS OF WETLANDS
The rate of wetland loss cannot be quantified in most
countries, but is relatively well-documented in the USA.
Fig. 22.5 illustrates the difference between the distribution
of wetlands some 200 years ago and at the present day, and
the lower map in Fig. 22.5 also shows percentage loss
between the 1780s and the 1980s in each of the states.
Table 22.10 gives figures for the loss of wetlands in some
of the states.
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e fo) e e fo) fo) fo)
e fo) e e e fo)
fo) fo) e e
e fe) fe)
fo) fo) e fo) fo)
° fo) e fo) e ° °
fo) fo) fo) fo) fo} fo) fo)
Notes: © Common and important value of that wetland type; © less common/important.
Source: Dugan, P.J. (Ed.) 1990. Wetland Conservation: a review of current issues and required action. UCN, Gland, Switzerland.
298
Wetlands
Table 22.5 The causes of wetland loss
7
4 2
8 8 2 : ® a
Cie ine 2
oe tae au iar ph ae
Ww [o) re LE = ao 7)
Human actions
Drainage for agriculture, forestry and mosquito control e ® e ® fo) C) ®
Dredging and stream channelisation for navigation and flood protection e fe)
Filling for solid waste disposal, roads, and commercial, residential and e e® e e fo)
Conversion for aquaculture/mariculture e (o) fe} (e) {e)
Construction of dykes, dams, levees, and seawalls for flood control, water e e e e fe)
Discharges of pesticides, herbicides, nutrients from domestic sewage and e e e e e
Mining of wetlands for peat, coal, gravel, phosphate and other materials fo) fo) fe) e e® ®
Groundwater abstraction fe) e
Sediment diversion by dams, deep channels and other structures e e ® e
Hydrological alterations by canals, roads and other structures ® ® e® e e
Subsidence due to extraction of groundwater, oil, gas and other minerals e (o) e e
Natural causes
Subsidence fe) fo) fe) (e) fe)
Sea-level rise e e
Drought . e e e e fe} (o}
Hurricane and other storms e e fe) {e)
Erosion e e fo) fo)
Biotic effects e e e
Notes: © common and important cause of wetland degradation and loss; © present, but not a major cause of loss.
Source: Dugan, P.J. (Ed.) 1990. Wetland Conservation: a review of current issues and required action. YUCN, Gland, Switzerland.
Table 22.6 Major threats to wetlands Table 22.7 Major threats to wetlands in
in Latin America and the Asia
Caribbean
THREAT INCIDENCE
THREAT INCIDENCE (% OF SITES)
(% OF SITES)
Hunting and associated disturbance 32
Pollution 31.0 Human settlement/encroachment 27
Hunting and associated disturbance 30.5 Drainage for agriculture 23
Drainage for agriculture and ranching 19.0 Pollution 20
Disturbance from recreation 11.5 Fishing and associated disturbance 19
Reclamation for urban Commercial logging/forestry 17
and industrial development 10.5 Wood cutting for domestic use 16
Forestry activities 10.0 Degradation of watershed/soil erosion/siltation 15
Fishing and associated disturbance 10.0 Conversion to aquaculture ponds or salt pans 11
Diversion of water supply 9
Source: Finlayson, M. and Moser, M. (Eds) 1991. Wetlands. Facts pe EOI ERE YSMOmes ne SOCk Sa) RAs See ee BYMOMIESHOIStece s)
on File Limited, Oxford. Based on information from 620 wetland sites
described in Scott, D.A. and Carbonell, M. (Eds) 1991. A Directory Source: Scott, D.A. and Poole, C.M. 1989. A Status Overview of
of Neotropical Wetlands. UCN, Cambridge and IWRB, Slimbridge. Asian Wetlands. No.53. AWB, Kuala Lumpur.
Notes: These are for 734 sites for which information on threats is
known. Only 107 of these are not threatened in one way or another
(Table 22.8).
299
1. Biological Diversity
Figure 22.3 Asian wetlands: distribution of sites
o
2
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@ =
= Ss
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7 >
= s
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300
Wetlands
Figure 22.4 Asian wetlands: threatened sites
a
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o
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MM © Degraded site
301
1. Biological Diversity
Table 22.8 Severity of threats to wetlands of international importance in Asia
NUMBER OF DEGREE of THREAT % SITES WITH
SITES MODERATE TO
KNOWN NONE LOW MOD HIGH HIGH THREAT
Bangladesh 11 1 1 5 4 82
Bhutan 5 3 - 1 1 40
Brunei 3 - 2 1 - 33
China 105 30 34 36 5 39
Hong Kong 3 1 - 24 - 67
India 88 4 44 22 18 45
Indonesia 129 1 54 66 8 57
Japan 38 8 11 17 2 50
Cambodia 3 = 1 2 - 67
Korea, DPR 5 5 = - - (e)
Korea, Rep. 19 5 3 6 5 58
Laos 3 - 1 2 - 67
Malaysia 37 - 5 22 10 86
Mongolia 30 23 5 2 - 7A
Myanmar 16 - 7 8 1 56
Nepal 14 2 7 4 1 36
Pakistan 42 1 20 15 6 50
Papua New Guinea 27 14 8 4 - 15
Philippines 49 2 13 24 10 69
Singapore 6 - 2 3 1 67
Sri Lanka 31 2 8 13 8
Taiwan 12 1 4 5 2 58
Thailand 36 1 18 14 3 47
Viet Nam 23 3 14 4 2 26
TOTAL 734 107 262 278 87 50
Source: Scott, D.A. and Poole, C.M. 1989. A Status Overview of Asian Wetlands. No.53. AWB, Kuala Lumpur, Malaysia.
In New Zealand, it is estimated that over 90% of natural
wetlands have been destroyed since European settlement and
drainage is still occurring (Dugan, 1990). Fig. 22.6 shows
the wetland areas of Waikato Basin in North Island in 1840
and in 1976. There were 1,614km? of wetland in 1840 and
2,62km? in 1976; even less remains now (Finlayson and
Moser, 1986).
Dams have been the cause of considerable reduction in the
areas of floodplains. Table 22.11 gives some estimates of
how floodplains in Africa will be reduced by the year 2020
as a result of construction of dams upriver.
CONSERVATION OF WETLANDS
Protected Areas
Until recently, wetland conservation was largely confined
to establishing protected areas, but wetlands are influenced
by activities well beyond their boundaries so this often does
not have the required effect. For instance, Scott and Poole
(1989) give tables in which the degree of threat to fully
protected sites in Asia and to those which are unprotected
are estimated (Table 22.12). They note that the creation of
protected areas in Asia has been successful to a certain
extent, but that an analysis of threats to the sites show that
the sole or principal threat is frequently from an external
source outside the control of the reserve manager. While
problems such as illegal settlement, overhunting, logging
302
and overgrazing can, in theory, be controlled within a
protected area, threats such as siltation from soil erosion in
the watershed, pollution from urban or industrial areas,
contamination with pesticides or fertilizers and disruption of
water supplies due to damming and flood control projects
upstream are all usually outside the jurisdiction of the
authorities concerned with conserving a protected area.
There are 69 protected wetlands in Asia that are moderately
or highly threatened (Table 22.12). For 20 of these, the
problems are solely or principally external in origin and for
a further 13 this is a significant cause of the threat (Scott
and Poole, 1989).
The Ramsar Convention and conservation
The Convention on Wetlands of International Importance
especially as Waterfowl Habitat (or the Ramsar Convention)
provides the principal intergovernmental forum for the
promotion of international cooperation for wetland
conservation. There are more than 60 Contracting Parties.
Some of the principal obligations of the parties are:
@ to designate wetlands of international importance for
inclusion on a list of ‘Ramsar sites’ and to advise the
Bureau of any change in their ecological character
to formulate and implement planning so as to promote
conservation of listed sites
to formulate and implement planning so as to promote
the wise use of wetlands
Table 22.9
Bangladesh
1 Chalan Beel*
2 Haor Basin of Sylhet and Eastern Mymensingh
3 Dubriar Haor*
4 The Sundarbans
5 Wetlands in Pablakhali Wildlife Sanctuary
6 Chokoria Sundarbans*
Bhutan
7 Boomthang Valley
Burma (Myanmar)
8 Irrawaddy Delta
People’s Republic of China
9 Yancheng Marshes
10 ~~ Shijiu Hu
11 Shengjin Hu and the lower Yangtze Lakes
12 Shengjin Hu
13 Xi Jiang (Pearl River) Delta*
14 Tuosu Hu (Kurlyk Nor) and Kuerhleiko Hu
15 Dal Lake
16 Wular Lake
17 Harike Lake
18 Jheels in the vicinity of Haidergarh*
19 Dahar and Sauj (Soj) Jheels
20 Southern Gulf of Kutch
21 Gulf of Khambhat
22 Khabartal
23 Dipor (Deepar) Bheel
24 Logtak Lake
25 Salt Lakes Swamp*
26 The Sunderbans
27 Chilka Lake
28 Kolleru Lake
29 Estuaries of the Karnataka coast
30 Kaliveli Tank and Yedayanthittu Estuary
31 The Cochin Backwaters
32 Wetlands in the Andaman and Nicobar Islands
Indonesia
33 Banyuasin Musi River Delta
34 Muara Cimanuk*
35 Sukolilo
36 Cilacap and Segara Anakan
37 Danau Bankau and other swamps in the Barito
Basin*
38 Banau Sentarum
39 Wetlands in Manusela Proposed National Park
40 Wasur and Rawa Biru
41 Shonai-Fujimae Tidal Flats and Inner Ise Bay
42 Lake Shinji and Lake Nakaumi
Republic of Korea
43 South Kanghwa and North Yongjong Mudflats
44 Mudflats of South Yongjong and adjacent islands
45 Namyang Bay
46 Asan Bay
47 Kum, Mankyung and Tangjin Estuaries
Wetlands
The most seriously threatened wetlands in Asia
Malaysia
48 Sedili Kecil Swamp Forest
49 Klang Islands: Pulau Ketam”*
50 Kapar Forest Reserve
51 North Selangor Swamp Forest
52 Marintaman Mengalong*
53 Tempasuk Plain
54 Lawas Mangroves
55 Trusan-Sundar Mangroves
56 Limbang Mangroves
57 Maludam Swamp Forest
58 Sarawak Mangrove Forest Reserve
59 Begnas Tal*
Pakistan
60 Khabbaki Lake*
61 Siranda Lake*
62 Hawkes Bay/Sandspit Beaches and adjacent creeks
63 Clifton Beach
64 Korangi and Gharo Creeks
65 The Outer Indus Delta
Philippines
66 Pangasinan Wetlands*
67 Manila Bay*
68 Laguna de Bay
69 Tayabas Bay including Pagbilao Bay
70 Inabanga Coast
71 Mactan, Kalawisan and Cansaga Bays*
72 Agusan Marsh
73 Lake Leonard*
74 Davao Gulf
75 Liguasan Marsh
Singapore
76 Serangoon Estuary*
Sri Lanka
77 Thandamannar Lagoon*
78 Chundikkulam Lagoon
79 Chalai Lagoon*
80 Periyakarachchi and Sinnakarachchi Lagoons*
81 Mahaweli Ganga Floodplain System
82 Maha Lewaya and Karagan Lewaya
83 Lunama Kalapuwa and Kalametiya Kalapuwa*
84 Bellanwilla-Attidiya Marshes
Taiwan, R.O.C.
85 Tatu Estuary
86 Tungshih (Ton-Shou) Mangroves*
Thailand
87 Gulf of Thailand
88 Pak Phanang Estuary
89 Pa Phru
Socialist Republic of Viet Nam
90 Red River Delta
91 Red River Estuary
92 Mekong Delta
93 Nam Can Mangrove Forest
Source: Scott, D.A. and Poole, C.M. 1989. A Status Overview of Asian Wetlands. No.53. AWB, Kuala Lumpur, Malaysia.
Note: * Sites marked with an asterisk are considered to be already too degraded to merit any special conservation effort.
© to make national wetland inventories
® to make environmental impact assessments before
transformations of wetlands
© to establish nature reserves on wetlands and provide
adequately for their wardening
® to train personnel to manage, research and warden
wetlands.
Each country’s Wetlands Conservation Programme will,
obviously, be determined by the characteristics of its natural
resources, the problems they face and the status of the
current conservation efforts, as well as by the financial and
human resources available (Dugan, 1990).
The quantity and quality of information on wetlands needs
to be increased and improved and it is essential that their
value is understood and that the benefits of them are seen.
Only when this happens will the rate of wetland loss be
substantially reduced.
1. Biological Diversity
Figure 22.5 Loss of wetlands in selected states of the USA
circa 1780s
oo
% Wet land er 5-12 25-50
42-25 iy) 50-55
Source: Dahl, T.E. 1990. Wetlands Losses in the United States 1780s to 1980s. US Department of the Interior, Fish and Wildlife Service,
Washington, DC.
304
Wetlands
Figure 22.6 Wetland loss in New Zealand, 1840 to 1976
a8,
Hl Wetland Areas
Source: Finlayson, M. and Moser, M. (Eds) 1991. Wetlands. Facts on File Limited, Oxford.
Table 22.10 Loss of wetlands in selected states of the USA
STATE ESTIMATES OF WETLAND ESTIMATES OF WETLAND % LOST
PRESENT IN 1780s (km?) PRESENT IN 1980s (km?)
California 20,000 1,837 91
Ohio 20,000 1,954 90
lowa 16,200 1,707 89
Indiana 22,700 3,038 87
Illinois 33,000 5,077 85
South Carolina 25,960 18,855 27
West Virginia 540 413 24
Georgia 27,694 21,442 23
Maine 26,140 21,041 20
New Hampshire 890 809 9
TOTAL* 895,000 422,397 53
Alaska 688,790 687,980 0.1
Source: Dahl, T.E. 1990. Wetlands Losses in the United States 1780s to 1980s. US Department of the Interior, Fish and Wildlife Service,
Washington, DC.
Note: * for all 48 conterminous states.
Table 22.11 Projected reduction in floodplain area as a result of dams
FLOODPLAIN AREA IN 1960 AREA IN 2020
(km?) (km?)
Senegal Delta 3,000 300
Senegal Valley 5,500 550
Niger Delta 30,000 27,000
Niger Valley 3,000 © 1,500
Sokoto and Rima 1,000 500
Hadejia Komadugu 3,800 380
Logone 11,000 6,600
Source: Dugan, 1990; modified after Drijver, C.A. and Rodenburg, W.F. 1988. Water Management at a Cross Roads: the case of the Sahelian
wetlands. Paper presented at the International Symposium on Hydrology of Wetlands in Semi-arid and Arid regions. Seville, Spain.
305
1. Biological Diversity
Table 22.12 Degree of threat to protected and unprotected Asian wetlands
Fully protected sites
REGION NO. OF
SITES
NONE
South Asia 70 6
Southeast Asia 64 2
East Asia 57 27
TOTAL 191 35
Unprotected sites
REGION NO. OF
SITES
NONE
South Asia 107 4
Southeast Asia 176 18
East Asia 104 41
TOTAL 387 63
DEGREE OF THREAT % MOD &
HIGH
LOW MODERATE HIGH
39 20 5 36
33 27 2 45
15 14 1 26
87 61 8 36
% MOD &
DEGREE OF THREAT HIGH
LOW MODERATE HIGH
46 33 24 53
65 77 16 53
31 25 7 31
142 135 47 47
Source: Scott, D.A. and Poole, C.M. 1989. A Status Overview of Asian Wetlands. No.53. AWB, Kuala Lumpur, Malaysia.
References
Aselmann, I. and Crutzen, P.J. 1989. Global distribution of natural
freshwater wetlands and rice paddies, their net primary
productivity, seasonality and possible methane emissions. Journal
of Atmospheric Chemistry 8:307-358.
Dahl, T.E. 1990. Wetlands Losses in the United States 1780s to 1980s.
US Department of the Interior, Fish and Wildlife Service,
Washington, DC.
Drijver, C.A. and Rodenburg, W.F. 1988. Water Management at a
Cross Roads: the case of the Sahelian wetlands. Paper presented at
the International Symposium on Hydrology of Wetlands in
Semi-arid and Arid regions. Seville, Spain.
Dugan, P.J. (Ed.) 1990. Wetland Conservation: a review of current
issues and required action. TUCN, Gland, Switzerland.
Finlayson, M. and Moser, M. (Eds) 1991. Wetlands. Facts on File
Limited, Oxford.
306
Gore, A.J.P. 1983. Introduction. In: Gore, A.J.P. (Ed.), Ecosystems
of the World (4A). Mires: swamp, bog, fen and moor. Vol. 1.
Elsevier, Amsterdam. Pp.1-34.
Matthews, E. and Fung, I. 1987. Methane emission from natural
wetlands: global distribution, area and environmental characteristics
of sources. Global Biogeochemical Cycles 1(1):61-86.
Scott, D.A. and Carbonell, M. (Eds) 1986. A Directory of Neotropical
Wetlands. TUCN, Cambridge and IWRB, Slimbridge.
Scott, D.A. and Poole, C.M. 1989. A Status Overview of Asian
Wetlands. No.53. AWB, Kuala Lumpur, Malaysia.
Chapter contributed by Caroline Harcourt. World map of
distribution of wetland areas based, by kind permission, on
continent maps in Finlayson and Moser (1991).
23. CORAL REEFS
OCCURRENCE OF REEFS
Coral reefs are tropical shallow water ecosystems largely
restricted to the seas between the latitudes of 30°N and
30°S. The exact extent of coral reefs in the world is
unknown and is very difficult to estimate. Smith (1978) has
calculated that there are 600,000km? of reefs to a depth of
30m. About 60% of this total occurs in the Indian Ocean
region; approximately 14% in the Caribbean, 13% in the
South Pacific (including eastern Australia) and 12% in the
North Pacific (including the Galapagos and west coast of
North America). The remaining 1% is divided between the
South Atlantic and the Eastern Pacific. The map in Fig.
23.1 shows the general location of the coral reef systems of
the world.
Coral reefs are one of the most productive and diverse of
all natural ecosystems; they are the marine equivalents of
the rain forests (Bourliére and Harmelin-Vivien, 1989).
Their richness stems from the steady availability of a wide
and diverse array of food resources and the extreme
heterogeneity of the environment, with the corals forming
a complex tri-dimensional structure providing a vast array
of habitats for a great variety of organisms. Data on generic
richness of corals and reef fish diversity at a number of
representative sites are shown in Table 12.5, with notes on
the fishes of coral reefs (Chapter 12).
The true reef-building coral polyps (stony or hermatypic
corals) are ones that collectively deposit calcium carbonate
to build colonies. The term ‘reef’? is used here for a
population of stony corals which continues to build on
products of its own making (Stoddart, 1969). However, not
all reefs are constructed predominantly of coral. For
instance, several genera of red algae grow as heavily
calcified encrustations which bind the reef framework
together, forming structures such as algal ridges.
Alternatively, populations of ahermatypic and non-symbiotic
corals exist which do not build reefs, while other
populations do not build on themselves. These have been
termed coral assemblages or communities.
Reefs fall into two basic categories: shelf reefs, which form
on the continental shelf of large land masses; and oceanic
teefs, which develop in deeper waters often in association
with oceanic islands. Within these two categories are a
number of different reef types: fringing reefs which grow
close to the shore; patch reefs which form on irregularities
on shallow parts of the sea bed; bank reefs which occur in
deeper waters, both on the continental shelf and in oceanic
waters; barrier reefs which develop along the edge of a
continental shelf or through land subsidence in deeper
waters and are separated from the mainland or island by a
relatively deep, wide lagoon; and atolls, which are roughly
circular reefs around a central lagoon and are typically
307
Coral Reefs
found in oceanic waters, probably corresponding to the
fringing reefs of long submerged islands.
VALUES AND THREATS
The World Conservation Strategy (IUCN/UNEP/WWF,
1980) identifies coral reefs as one of the "essential
ecological processes and life-support systems" necessary for
food production, health and other aspects of human survival
and sustainable development. Reefs protect the coastline
against waves, prevent erosion and contribute to the
formation of sandy beaches and sheltered harbours. They
also provide nutrients and breeding grounds for many
commercial and subsistence fish species, as well as a habitat
for numerous molluscs and crustaceans that are also caught
for food. The tourist industry is another important source
of income to many countries and much of this is related to
the presence and condition of reefs.
Damage to coral reefs can be caused through natural events,
including storms and hurricanes, climate changes, disease
and predators of coral. Humans also have an impact
through pollution (sewage, pesticides, fertilizers, industrial
waste, etc.), sedimentation (often following land clearance
and subsequent erosion inland), and over-exploitation of
reef resources (intensive recreational use, coral mining).
Table 23.1 below lists in summary form: the reef resources
found in tropical countries, their use, disturbances to them
and what legislation or management occurs. The
information has been extracted from UNEP/IUCN
(1988a,b,c). Fig. 23.1 is based largely on this same source.
References
Bourliére, F. and Harmelin-Vivien, M.L. 1989. Species diversity in
tropical vertebrates: an ecosystem perspective. In: Harmelin-Vivien,
M.L. and Bourligre, F. (Eds), Vertebrates in Complex Tropical
Systems. Springer-Verlag, New York.
TUCN/UNEP/WWE 1980. World Conservation Strategy: living
resource conservation for sustainable development.
TUCN/UNEP/WWF, Gland, Switzerland.
Smith, S.V. 1978. Coral reef area and contributions of reefs to
processes and resources of the world’s oceans. Nature 273:225.
Stoddart, D-R. 1969. Ecology and morphology of recent coral reefs.
Biological Review Cambridge Philosophical Society 44:433-498.
UNEP/IUCN (1988a). Coral Reefs of the World. Vol. 1: Atlantic and
Eastern Pacific. UNEP Regional Seas Directories and
Bibliographies. TUCN, Gland, Switzerland and Cambridge,
UK/UNEP, Nairobi, Kenya. 373pp., 38 maps.
UNEP/IUCN (1988b). Coral Reefs of the World. Vol. 2: Indian
Ocean, Red Sea and Gulf. UNEP Regional Seas Directories and
Bibliographies. TUCN, Gland, Switzerland and Cambridge,
UK/UNEP, Nairobi, Kenya. 389pp., 36 maps.
UNEP/IUCN (1988c). Coral Reefs of the World. Vol. 3: Central and
Western Pacific. UNEP Regional Seas Directories and
Bibliographies. TUCN, Gland, Switzerland and Cambridge,
UK/UNEP, Nairobi, Kenya. 329pp., 30 maps.
1. Biological Diversity
Figure 23.1 Coral reef systems of the world
2000 4000 6000
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308
Coral Reefs
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323
1. Biological Diversity
24. MANGROVES
THE MANGROVE HABITAT
Mangroves are the characteristic littoral plant formations of
sheltered, low-lying tropical and subtropical coasts.
The species found in these habitats are a diverse collection
of trees and shrubs that have adapted to salty, inundated
environments. The mangrove species can be divided into
two groups: the exclusive species found only in mangrove
habitats and the non-exclusive species which may be
important in the mangrove community but are not restricted
to it. Saenger et al. (1983) list 60 species in the former
group, in 22 genera, and give a list of 23 species, in 16
genera, which are some of the important non-exclusive
species. The appearance of mangroves is far from uniform;
they vary from closed forests 40-50m high in parts of South
America to stunted shrubs less than 1m high, which can be
in discrete and widely separated clumps (Finlayson and
Moser, 1991).
In addition to the mangroves themselves, a wide variety of
organisms are associated with the mangrove system, and the
habitat is critical to many. Such organisms include a
number of epiphytes, parasites and climbers among the
flora, and large numbers of crustaceans, molluscs, fishes
and birds among the fauna. Table 24.1 provides indicative
information on species richness in these groups.
Current information on the area of mangrove habitat in each
country where it occurs, together with the number of
Table 24.1 Species richness in groups
associated with mangroves
TAXONOMIC GROUP ASIA CARIBBEAN
W. and ATLANTIC
Bacteria 10 -
Fungi 25 -
Algae 65 105
HIGHER PLANTS
Bryophytes/Ferns 35 2
Monocotyledons 73 20
Dicotyledons 110 28
ANIMALS
Protozoa 18 3
Sponges/Bryozoa 5 36
Coelenterata/Ctenophora 3 42
Non-polychaete worms 13 13
Polychaetes 11 33
Crustaceans 229 87
Insects/Arachnids 500 -
Molluscs 211 124
Echinoderms 1 29
Ascidians (e) 30
Fish 283 Ales
Reptiles 22 3
Amphibians 2 2
Birds 177 138
Mammals 36 5
Source: Abridged from Saenger, P., Hegerl, E.J. and Davie, J.D.S.
(Eds) 1983. Global Status of Mangrove Ecosystems. Commission on
Ecology Papers Number 3. IUCN, Gland, Switzerland.
Note: Estimates refer to two regions only; they are not fully
comprehensive but provide general indications of relative species
richness; missing data shown by -.
protected areas known to include mangroves, is given in
Table 24.2.
VALUE OF MANGROVES
The uses and values of mangroves to humans are many and
varied. The wood from mangrove trees is used by local
people for building materials for houses, fence-poles,
material for fish traps, and so forth, and is also harvested
on a large scale by international companies, particularly for
pulp and particle board. The mangroves are also a
significant source of fuel, both firewood and charcoal. The
most important species for this purpose are those belonging
to the genus Rhizophora, as this wood is heavy and clean
burning. Another potential source of fuel in the mangrove
habitat is the Nypa palm (Nypa fruticans), which produces
a sugar that can be converted into aloohol and be used as a
transport fuel (Saenger et al., 1983).
Other products from the mangrove habitat include shellfish,
crustaceans and fish. These are harvested both on a
subsistence basis and commercially. For instance, in the
mangroves of Sierpe, Costa Rica, the five million shellfish
(Anadara sp.) harvested annually are worth US$85,000 to
the local communities which collect them (Lahmann, 1989).
Most of the larger commercial penaeid shrimps are
mangrove-dependent, and fisheries for shrimps and prawns
are major sources of export earnings in many tropical
countries. In addition to providing habitat for adult fish, the
mangroves are essential spawning and nursery areas for
many species of marine fish. In the Gulf of Mexico, 90%
of the fish harvest, worth US$700 million per year, consists
of species which are dependent on mangroves and other
coastal wetlands at some stage in their life cycle (Dugan,
1990). Mangrove communities are amongst the most
productive ecosystems in the world.
Domestic animals feed on mangrove foliage in many
countries. In Pakistan, camels are herded down to the coast
to feed on the mangroves in the dry season; water buffalo
graze them in parts of Asia and Australia, while in Africa
goats and cattle feed on the foliage (Dugan, 1990).
Mangroves stabilise shorelines and decrease coastal erosion
by reducing the energy of waves and currents and by
holding the bottom sediment in place with plant roots. They
also act as windbreaks and protection from coastal storms,
forming a cost-free, self-repairing barrier.
Mangrove habitats can be used for tourism, education and
scientific study. For instance, the Bengal Tiger Panthera
tigris population in Sundarbans mangrove is a focus for
tourism in Bangladesh, and thousands of visitors go to
Trinidad’s Caroni Swamp mangrove area every year to
view the great numbers of Scarlet Ibis Eudocimus ruber and
other rare or endangered birds found there.
THREATS TO MANGROVE HABITATS
Vast areas of mangroves are being destroyed either directly
or as a secondary result of other activities. There are
numerous reasons for the destruction and frequently short-
Mangroves
Table 24.2 Mangroves
COUNTRY TOTAL AREA PROTECTED COUNTRY TOTAL AREA PROTECTED
OF MANGROVES AREAS WITH OF MANGROVES AREAS WITH
(in hectares) MANGROVES (in hectares) MANGROVES
ASIA NORTH AND CENTRAL AMERICA (continued)
Bahrain 40 1 Martinique 2,200 72
Bangladesh 410,000 5 Mexico 1,420,200 9)
Brunei 7,000 3 Montserrat 7 {e)
Cambodia 10,000 Oo Netherlands Antilles (total) 2,200 5
China 67,000 4 Bonaire 1,000 2
Hong Kong s 9 Curacao 300 1
India 356,000 (34) St Martin and Barthélémy 100 {0}
mainland (306,000) (9) Nicaragua 60,000 9
Andaman Is (50,000) 25 Panama 297,532 23
Indonesia 4,251,011 152 Puerto Rico 6,497 13
Irian Jaya 2,934,000 19 St Lucia 179 5
Java and Bali 51,885 30 St Kitts and Nevis 20 -
Kalimantan 383,450 28 St Vincent * 2
Lesser Sunda Is 5,508 16 Trindad and Tobago 9,000 6
Moluccas 100,000 12 Turks and Caicos Is ¢ 5
Sulawesi 99,833 18 USA 280,594 -
Sumatra 657,335 29 Alabama 25 -
Japan (Ryuku Is) 400 4 California 150 -
Iran 2a F 12 3 Florida 274,857 (47)
Malaysia 630,000 99 Louisiana 2,956 (2
Peninsula 105,000 48 Mississippi 250 -
Sabah 350,000 35 Texas 2506 (3)
Sarawak 175,000 16 Hawaii * 10)
Maldives iv (0) Virgin Islands (British) 7 10
Myanmar 517,000 6 Virgin Islands (United States) 310 4
ppen ‘ : SOUTH AMERICA
Pakistan 249,500 2
Philippines 400,000 59 Brazil 250,000 (32)
Qatar * (e) Colombia 501,300 12
Saudi Arabia * - Ecuador 182,108 6
Singapore 1,800 2 Galapagos Is (200) 4
Sri Lanka 120,000 9 French Guiana 5,500 3
Taiwan 174 4 Guyana 80,000 -
Thailand 268,693 17 Peru 6,346 3
Viet Nam (South) 370,000 2 Suriname 115,000 6
UAE e 6 Venezuela 673,569 13
Biemen ; 2 OCEANIA
NORTH AND CENTRAL AMERICA 5
American Samoa e co)
Anguilla S (0) Australia 1,161,700 218
Antigua and Barbuda 600 & 900 0 Capital Territory Bo 74
Aruba 100 2 New South Wales 10,673 33
Bahamas 233,200 10 Northern Territory . 20
Barbados 12 1 Queensland - 113
Belize 78,317 13 South 20,100 21
Bermuda 17 8 Victoria i 18
Cayman Islands 11,655 11 Western i 11
Costa Rica 35,000 11 Fiji 19,700 3
Cuba 626,000 (23) French Polynesia (Society Is) = fe)
North coast 131,000 - Guam ot 1
N coast Is and arch. 114,000 - Kiribati (Gilbert Is) i (0)
South coast 318,000 - Marshall Is = (0)
S coast Is 38,000 - Micronesia, Fed. States i ie)
S Is & | de Pinos 25,000 - Nauru az te)
Dominica 2s 1 New Caledonia 20,000 2
Dominican Rep. 23,500 6 New Zealand 19,800 (15)
El Salvador 45,000 6 Palau 4,708 1
Grenada by 11 Papua New Guinea 200,000 31
Guadeloupe 5,700 2a Solomon Islands 64,200 ie)
Guatemala 16,000 6 Tonga 1,000 2
Haiti 18,000 {o} Tuvalu 47 fo)
Honduras 117,000 10 Vanuatu * Oo
Jamaica 20,200 12 Western Samoa <1,000 2
- 325
1. Biological Diversity
Table 24.2 Mangroves (continued)
COUNTRY TOTAL AREA PROTECTED
OF MANGROVES AREAS WITH
(in hectares) MANGROVES
AFRICA
Angola 110,000 2
Benin 3,000 1
Cameroon 306,000 1
Comoros * {e)
Congo (2,000) 1
Cote d'Ivoire (2,000) 2
Djibouti i! fe}
Egypt = 2
Equatorial Guinea 20,000 3
Ethiopia # 1
Gabon 250,000 2
Gambia 66,000 3
Ghana (2,000) 3
Guinea 223,000 1
Guinea-Bissau 236,000 3
Kenya 45,000 14
Liberia (20,000) 2
Madagascar 325,560 4
COUNTRY TOTAL AREA PROTECTED
OF MANGROVES AREAS WITH
(in hectares) MANGROVES
AFRICA {continued)
East coast 4,815 -
West coast 320,745 -
Mauritania * 3
Mauritius 7 6
Mozambique 85,000 8
Nigeria 3,238,000 1
Réunion < 2
Seychelles = 2
Senegal 169,000 3
Sierra Leone 250,000 6
Somalia 10,000 6
South Africa 673 11
Sudan ¥ 1
Tanzania 133,540 2
Mainland 115,476 -
Zanzibar Is 18,064 -
Togo if te)
Zaire 53,000 1
Source: WCMC Protected Areas Data Unit. 1991. Draft list of Protected Areas with mangrove habitats. Much information in this list was provided
by S.C. Snedaker.
Notes: * Country notes are provided in the Draft list; in many cases these notes indicate that no numerical estimate was obtainable; in others,
additional island-specific estimates are given. (-) No information. ( ) Figure from list of protected areas, considered an estimate.
term exploitation for economic gain takes precedence over
long-term benefits which have both economic and natural
value.
In many areas, the demand for fuelwood from mangroves
is well above a sustainable level and it is increasing as the
human population increases. In addition, the commercial
use of the wood, for pulp in particular, results in some
areas being more or less clear-felled. Natural regeneration
frequently does not occur and often the area is converted to
other forms of land-use such as agriculture or aquaculture.
In many instances where attempts have been made to
convert mangroves to agricultural land, the soil becomes
extremely acid because of the oxidation of the pyrite
sulphur which is commonly found in large quantities in the
mangrove soils. This, combined with the high concentration
of soluble salts in the soil, leads either to crop loss or to a
considerable reduction in productivity (Saenger ef al.,
1983).
The conversion of mangroves areas to aquaculture gives
rise to a number of problems. Much of the mangrove flora
and fauna in the areas surrounding the ponds is destroyed
because of major changes in drainage conditions, nutrient
availability and frequency of tidal inundation, as well as
being adversely affected by run-off from ponds and
channels. The acid sulphate soils that have an adverse effect
on crops also inhibit algae growth, which the fish feed on,
and may kill the prawns or fish directly by poisoning them.
Conversion to aquaculture ponds is a particular threat in the
Asian region, although in the Indo-Pacific area it was
estimated, in 1977, that 1.2 million ha of mangrove forest
had already been converted to aquaculture ponds (Saenger
et al., 1983). The building of ponds for extraction of salt
water can, especially in arid and semi-arid areas, cause
extensive damage to mangroves. The land has to be cleared
326
of all trees and shrubs, levelled and dyked; a canal system
has to be built and the soil surface compacted, so that even
if the ponds are later abandoned the chemical and physical
properties of the soil have been so changed that
recolonisation by mangroves is impossible.
Mangroves are also converted for urban and industrial
development, commonly for housing, tourist facilities,
airports and small ports. Many of the mangroves that are
not directly destroyed by these developments are affected by
loss of freshwater and by pollution from numerous different
sources. Rubbish and solid wastes are often deliberately
dumped in mangrove habitats. Mining within the mangrove
system completely destroys the habitat, while mining in
adjacent areas causes variable adverse effects, foremost
among these being excessive silt deposition in the mangrove
system causing tree loss or reduced productivity. Chemical
wastes from mines are also frequently carried into coastal
areas where mangroves occur, with similar effects. Drilling
for oil occurs in some mangroves and both spillage and the
associated pipelines and roads which alter the drainage of
the area can be very destructive to the ecosystem.
Another threat to mangroves is a diversion or alteration of
the freshwater flow into them. In arid, semi-arid or
seasonally dry regions the mangroves are particularly
dependent on periodic inputs of freshwater, but in these
regions there is a high demand for freshwater and its flow
into the oceans is regarded as wasteful. Consequently,
rivers are often dammed or diverted so that their waters can
be used on land. Changes in land-use upstream, such as the
logging of a forest, can also affect the freshwater flow into
the mangroves. The reduction in freshwater results in the
gradual replacement of mangrove species with more salt-
tolerant and possibly less useful species. Mammals within
the mangrove system are affected by the lack of freshwater,
while fishery resources may be depleted by the higher
salinity and the reduced nutrients.
Much of the conversion of mangroves has occurred because
this habitat has, traditionally, been regarded as unproductive
wasteland. In many cases, government policies have
contributed to the destruction of the mangroves and it is
only as adverse effects of their disappearance are noted that
these policies are changing.
S27)
Mangroves
References
Dugan, P.J. (Ed.) 1990. Wetland Conservation: a review of current
issues and required action. IUCN, Gland, Switzerland.
Finlayson, M. and Moser, M. (Eds) 1991. Wetlands. Facts on File
Ltd, Oxford.
Lahmann, E. 1989. Formulacién de un proyecto de conservacién de
los recursos naturales para la zona de manglares de estero real,
Nicaragua. Mimeographed report, IUCN, San José, Costa Rica.
Saenger, P., Hegerl, E.J. and Davie, J.D.S. (Eds) 1983. Global Status
of Mangrove Ecosystems. Commission on Ecology Papers Number
3. IUCN, Gland, Switzerland.
PART 2
USES AND VALUES OF BIODIVERSITY
Part 1 of this book outlined the nature of biological diversity, the elements of which it
is comprised, and some ways in which diversity is measured. These themes were
illustrated by discussion and data on selected groups of organisms and habitats.
The intention in Part 2 is to introduce some of the ways in which humans use and benefit
from components of biodiversity, and to discuss aspects of the problems involved in
attempting to assign appropriate economic values to goods and services provided by
them.
Part 2 includes three chapters. Chapters 25 and 26 provide an introduction to human uses
of plants and animals respectively. The aim is to outline some of the principal uses and
selectively to present further detail, where possible by means of data tables, and usually
where the subject is of particular interest or is unfamiliar to many. No attempt has been
made to document comprehensively the entire range of uses to which natural resources
are put, nor to catalogue all the species involved.
It is an ecological imperative that humans depend on plants and, to a lesser extent, on
other animals for the basic requirements of existence, so we have not stressed this point.
Nor have we detailed the ecological functions at the habitat and landscape level that
collectively provide benefits in the form of services, largely because the role of diversity
in these functions is poorly-understood and difficult to quantify, although an area of
active international research. Some service functions, such as carbon-fixing and watershed
protection, could probably be performed as well by plantation forest monocultures as by
native multi-species forests.
Chapter 27 draws upon a growing literature on the application of economic theory to
biodiversity, in particular on the ways in which values can be attached to natural
resources or habitats. Whilst the discipline of the economist provides an interesting
perspective on biodiversity conservation, giving particular insight into the general forces
which drive habitat conversions, the analytic methods used can, again, be difficult to
apply to the concept of biological diversity.
The values that can be assigned to species or habitats may be sufficiently high to suggest
that cost/benefit analyses must often in the past have greatly undervalued their worth in
comparison to developments that impact upon them. On the other hand, whilst the
obvious value of keeping open future options provides a very powerful general argument
for conserving biological diversity, the economic values to be derived are extremely
difficult to quantify and can even be negative; many would regard the economist’s
viewpoint as here subsidiary to aesthetic and moral arguments.
329
Soe Be
eh A
tyme
7 alors 3
25. PLANT USE
Plant species provide an extremely wide range of useful
products relied on by people in all countries of the world.
A mixture of direct harvesting from the wild and cultivation
Tanging from basic subsistence farming to sophisticated
agricultural systems supplies food, medicines and a wealth
of raw materials. Plant biodiversity as a global resource
remains poorly understood, inadequately documented and
often wasted, but still retains immense potential for further
development of natural products.
Indigenous people in developing countries retain a basic
reliance on wild and traditionally cultivated plant species
that directly supply a wide range of their needs and often
display a remarkable knowledge of these local, often
undocumented, plant resources.
As well as the more obvious plant products such as food,
medicines, ornamental plants and timber, plants provide a
wide variety of resources used in industry and commerce.
To mention but a few, plant extracts are used in the
manufacture of glue, soaps, cosmetics, dyes, plastics,
lubricants and polishes. Plants provide an important source
of renewable energy, with Brazil, for example, obtaining
28% of its energy needs from sugarcane biomass resources.
This represents a major saving on petrol imports amounting
to US$8.9 billion from 1976 to 1985. An economic analysis
of various aspects of the value of plant diversity is given in
Chapters 27. This chapter provides a more general outline -
of the importance of plants in five major categories: food
plants; timber; rattans; medicinal plants; ornamentals.
FOOD PLANTS
One of the most fundamental values of plant biodiversity is
in supplying the world’s food. Originally plants were
consumed directly from the wild and gathering of wild
produce continues throughout the world today. Through the
processes of domestication wild plants became reservoirs of
new crop species and they are now an invaluable source of
genes needed to improve the world’s crops.
Of the estimated 250,000 species of flowering plants, only
about 3,000 have been regarded as a food source, although
most have probably been sampled at one time or another.
Others will have provided forage and browse for animals in
turn hunted or farmed by people. Around 200 plant species
have been domesticated for food, and of these about 15-20
are crops of major economic importance.
Relatively few botanical families account for the world’s
main domesticated plants. Gramineae and Leguminosae are
the most important, followed by the Cruciferae, Rosaceae,
Umbelliferae, Solanaceae and Labiatae. Other significant
families are the Chenopodiaceae, Araceae, Cucurbitaceae
and Compositae. Table 25.1 lists the nutritionally important
plants of the world and reflects the predominance of these
families. The species included are not all crops of major
economic importance but are the plants that account for the
bulk of food production.
Although relatively few plants contribute to food production
globally, at a local level plant resources provide a varied
source of nutritional needs. In one region of Peru, fruits of
331
Plant Use
193 species are regularly consumed; of these, 120 species
are exclusively wild-collected and a further 19 originate
from both wild and cultivated sources. Locally consumed
species such as these hold considerable potential as food
plants for wider use.
The history of food crops
The evolution of crop plants began between 5,000 and
10,000 years ago. It is now generally thought that
agriculture originated more or less simultaneously in
various parts of the world. The Fertile Crescent of the Near
East, centred on the area which is now Iraq, is well known
as the source of domestication of wheat and barley together
with certain pulses such as the lentil. Early agricultural
development based on the domestication of millets, also
took place in the loess regions north of the Huang He
(Yellow River) in China and in southern Mexico where
squashes, beans, peppers and maize were domesticated.
Agriculture is also thought to have developed independently
in the South American Andes.
From these early centres of agriculture the spread of
domesticated plants took place and, following conscious and
unconscious selection pressures, individual crops became
increasingly diverse.
In traditional agro-ecosystems newly domesticated plant
types and primitive cultivars diverged from their wild
ancestors. Nevertheless, occasional crosses continued to
occur between the early crops and their wild relatives and
allowed, for example, the incorporation of disease and pest
resistance genes harboured by wild parent plants. The
introduction of genes from wild and weedy relatives
increased the availability of crop genetic diversity for
further selection and improvement by farmers, and
increased the potential of crops to respond to changing
environmental conditions. Many cultivated species may not
have survived in domestication without the interchange of
genes between wild and crop populations (Oldfield, 1984).
Human migration and trade also contributed significantly to
the evolution of crop plants. When a species is introduced
to a new environment it often changes relatively quickly,
adapting to new ecological conditions. Furthermore, when
crops are taken from their source areas they may encounter
different wild relatives and cross with them. This has
occurred across continents, for example, with different
species of rice. In West Africa, an indigenous cultivated
rice Oryza glaberrima has hybridised with the introduced
common Asian rice O. sativa, enriching the rice gene pool
in the region. In this way, crops may develop secondary
centres of variation with greater genetic diversity than their
original centres of origin.
Processes of natural selection in response to new ecological
conditions have increased crop diversity and this in turn has
been enhanced by farmers selecting for particular
characteristics over thousands of years of cultivation. The
range of crops cultivated also increased through time as
additional species were brought into cultivation, and in
some cases weeds of the primary crops became important
food plants in their own right.
2. Uses and Values of Biodiversity
Table 25.1
FAMILY
Anacardiaceae
Araceae
Aquifoliaceae
Betulaceae
Bromeliaceae
Camelliaceae
Caricaceae
Chenopodiaceae
Compositae
Food crops of the world
SPECIES
Mangifera indica
Pistacia vera
Colocasia esculenta
Xanthostoma sagittifolium
Mex paraguariensis
Corylus avellana
Corylus maxima
Ananas comosus
Camellia sinensis
Carica papaya
Beta vulgaris
Chenopodium quinoa
Spinacia oleracea
Carthamus tinctorius
Cynara scolymus
FOOD
Mango
Pistachio
Taro
Yautia
Mate
Hazel
Filbert
Pineapple
Tea
Papaya
Sugar Beet
Quinoa
Spinach
Safflowerseed
Artichoke
ORIGIN
NE India, the majority of
fruit-bearing trees are more
or less wild.
Native to the Near East and
western Asia, cultivated in
the Mediterranean and
westem Asia for 3000-
4000 years.
India
A tropical American plant
developed by Amerindian
people.
Native to S. Brazil, _
Paraguay and N. Argentina,
cultivated throughout its
natural range. Leaves are
also still collected from wild
plants.
Europe and SW Asia.
Domesticated in the 17th
century.
SE Europe and western
Asia.
Thought to be a lowland
South American
domesticate.
Probably the lower Tibetan
mountains or Central Asia
Lowlands of eastern Central
America
Europe, developed as a crop
for sugar in the 18th
century.
A native American crop of
the high central Andes
developed by Indian
agriculturists in pre-
Colombian times.
Native to SW Asia
The cultivated species had
its origins in the Near East
Native to the Mediterranean
area and Canary Islands,
domesticated several
thousand years ago.
CONSERVATION
Wild species of mango are
threatened in Southeast Asia as a
result of deforestation and
replacement by commercial
species. WWF is funding
conservation of wild fruit trees in
Peninsular Malaysia.
Many wild populations have been
destroyed by forest clearance,
over-cutting for charcoal and
grazing.
Collection, preservation and
research are needed for aroid
cultivars. More than 1,000
cultivars of Co/ocasia exist as a
result of efforts by subsistence
farmers.
Species of wild pineapple are
native to botanically under-
explored parts of lowland South
America. They are now being
used in breeding programmes.
Collection and conservation of
clones from the upper Amazon
and Upper Orinoco is considered
desirable.
Truly wild teas probably no longer
exist. In cultivation a substantial
loss of genetic variability has
been anticipated which needs to
be countered by deliberate
conservation measures.
Wild populations of the related
Beta maritima are threatened in
parts of the Mediterranean.
One of the wild relatives of
safflower, the Moroccan endemic
Carthamus rhiphaeus is
considered Rare by !UCN.
Table 25.1
FAMILY
Compositae
(continued)
Convolvulaceae
Cruciferae
Cucurbitaceae
Dioscoreaceae
Euphorbiaceae
Gramineae
Food crops of the world (continued)
SPECIES
Helianthus annuus
Lactuca sativa
Ipomoea batatas
Brassica oleracea/B. rapa
Brassica juncea
Brassica napus,
B. rapa
Citrullus lanatus
Cucumis melo
Cucumis sativus
Cucurbita maxima,
C. moschata,
C. pepo
Dioscorea spp.
Manihot esculenta
Avena sativa
Echinochloa frumentacea
Eleusine coracana
Digitaria exilis
Hordeum vulgare
FOOD
Sunflowerseed
Lettuce
Sweet Potato
Cabbage
Mustardseed
Rapeseed
Melonseed
Melon/Water-
melon
Cucumber
Pumpkin,
Squash, Gourd
Yam
Cassava
Oats
Japanese
Barnyard Millet
Finger Millet
Fonio
Barley
333
ORIGIN
Domesticated in central
USA probably before the
arrival of maize, beans and
squash
Mediterranean
Central and South America
The wild cabbage is native
to Europe; development of
cultivars took place in the
Mediterranean region.
The primary centre of origin
is believed to be Central
Asia - Himalayas.
B. napus probably does not
exist in the wild.
Native to S Africa, chiefly in
the Kalahari Desert.
Africa, wild forms found in
eastern tropical Africa.
Native to India, probably
cultivated for over 3,000
years.
Domesticated in the
Americas at least 10,000
years ago.
Domestication of yams in
Asia, Africa and tropical
America took place
separately with different
species involved.
A cultigen, unknown in the
wild state.
Generally regarded as a
secondary crop, evolved in
W and N Europe from weed
oat components of wheat
and barley crops.
Different strains are thought
to have at least partially
different origins.
Central Africa. Taken to
India probably over 3,000
years ago where a second
centre of diversity became
established.
West Africa, thought to be
a cultigen.
One of the first crops
domesticated in the Near
East.
Plant Use
CONSERVATION
Some of the American varieties
have been preserved. A large
genetic reservoir exists among
the weed and wild sunflowers.
Wild gene pools are disappearing
owing to habitat loss.
The conservation of variability is
a major concern in breeding for
subsistence agriculture.
IBPGR has designated the
collection of wild forms of
B.oleracea as a conservation
priority. Several related
Mediterranean taxa are
threatened in the wild.
Large collections serve as
substantial gene pools and wild
material is widely distributed.
Many of the wild Cucurbita
species have restricted ranges.
Serious genetic erosion has
occurred among cultivated yams
and there is an urgent need to
collect and conserve genetic
diversity. There is little
information on the status of wild
relatives of yams.
The virtually unexplored wild
relatives are an important genetic
resource for crop improvement.
Centre of diversity of wild
relatives are in east-central Brazil,
NE Brazil and SW Mexico.
The potential of wild populations
in breeding programmes remains
to be determined.
This species is still capable of
genetic exchange with related
wild forms living in the same
area.
Concer about genetic erosion
e.g. in Ethiopia, where cultivars
are valuable for genetic resistance
to disease and improved
nutritional quality.
2. Uses and Values of Biodiversity
Table 25.1
FAMILY
Gramineae
(continued)
Grossulariaceae
Iliciaceae
Juglandaceae
Lauraceae
Food crops of the world (continued)
SPECIES
Oryza glaberrima,
O. sativa
Panicum miliaceum
Pennisetum americanum
Saccarhum officinarum
Secale cereale
Setaria italica
Sorghum bicolor
Triticum aestivum,
T. turgidum
Zea mays
Ribes nigrum, R. rubrum
Wlicium verum
Juglans regia
Persea americana
FOOD
Rice
Common Millet
Bulrush Millet
Sugarcane
Rye
Foxtail Millet
Sorghum
Wheat
Maize
Currants
Star Anise
Walnut
Avocado
334
ORIGIN
The origin of Asian rice O.
sativa is uncertain. The
African O. glaberrima
probably originated 3,500
years ago. Its primary
centre of diversity is the
swampy area of the Upper
Niger.
A millet of ancient
cultivation which is not
known in its wild state.
Probably in western tropical
Africa where the greatest
number of cultivated and
related wild forms occur. A
second centre of diversity
became established in India.
New Guinea
SW Asia, arising as a weed
of wheat and barley
Origin unknown in the wild
state, the crop is thought to
have arisen from the
common Old World weed S.
viridis.
Developed primarily from
the wild S. arundinaceum in
Africa.
Mediterranean and Near
East
Maize was domesticated in
prehistoric times in Mexico
and Central America.
Black and red currants are
native to northern Europe
and northern Asia, with the
black currant extending to
the Himalayas.
Domesticated in northern
Europe within the past 500
years.
China, Viet Nam
Native from SE China to
Europe
The crop originated in
Central America and has
been cultivated for several
thousand years.
CONSERVATION
As rice cultivation has become
more intensive, many wild
populations have disappeared.
The International Rice Research
Centre in the Philippines
coordinates the collection of
indigenous varieties. Little effort
has been made to conserve O.
glaberrima and its wild relatives,
however.
This species is still capable of
genetic exchange with related
wild forms living in the same
area.
Valuable germplasm of wild
sugarcane and related species has
been lost as a result of habitat
destruction in Malaysia, Indonesia
and Papua New Guinea.
A number of wild relatives are
restricted to small areas. There is
a need for further ex situ
conservation.
A wild species Z. perennis was
presumed extinct in the wild until
its rediscovery in 1977. A new
species was also discovered, Z.
diploperennis, and is now
protected in the Sierra de
Manantlan Biosphere Reserve,
Mexico.
Primitive wild relatives are
restricted to small areas in Central
America. The endangered caoba
tree from Ecuador
Caryodaphnopsis (Persea)
theobromifolia is a wild relative
resistant to blight.
Table 25.1
FAMILY
Leguminosae
Lecythidaceae
Liliaceae
Malvaceae
Moraceae
Food crops of the world (continued)
SPECIES
Arachis hypogaea
Cajanus cajan
Cicer arietinum
Glycine max
Lablab purpureus
Lens culinaris
Lupinus mutabilis
Phaseolus lunatus
Phaseolus vulgaris
Pisum sativum
Vicia faba
Vigna unguiculata
Bertholletia excelsa
Allium cepa: Allium
fistulosum
Allium sativum
Gossypium barbadense,
G. hirsutum
Ficus carica
FOOD
Groundnut
Pigeonpea
Chickpea
Soybean
Lablab bean
Lentil
Lupin
Lima bean
Haricot bean
Pea
Broad bean
Cowpea
Brazil nut
Onion
Garlic
Cottonseeds
Fig
335
ORIGIN
A cultigen domesticated
thousands of years ago in
South America.
The centre of origin is
assumed to be India
Western Asia
A cultigen not known in the
wild, soybean is thought to
have arisen as a
domesticate in the eastern
half of northern China.
Thought to be of Asian
origin, now widespread in
the tropics
The wild progenitor of the
cultivated lentil is Lens
orientalis, a Near Eastern
species.
A very variable cultigen of
the high Andes.
It is thought that separate
domestications occurred in
Central and South America
from conspecific geographic
races.
It is thought that separate
domestications occurred in
Central and South America
from conspecific geographic
races.
The wild progenitor is
unknown and the early
history of the pea crop is
unclear. Probable centres of
origin are Ethiopia, the
Mediterranean and Central
Asia.
Near East
The common cultivated
subspecies is thought to be
derived from wild plants in
Ethiopia several thousand
years ago.
Tropical South America.
Nuts are still collected from
wild trees as experimental
plantations have mainly
failed.
Central Asia
Known only in cultivation.
A. longicuspis, a species
endemic to central Asia,
may be its wild ancestor.
South America
Southern Arabia
Plant Use
CONSERVATION
Much unexplored genetic
variability in wild relatives of
potential importance in breeding
programmes. The protection of
perennial Arachis species in Latin
America is considered a
conservation priority.
Many of the wild relatives of
chickpea are threatened or rare.
Soybean cultivars grown in the
USA show a high degree of
genetic uniformity. The
germplasm base in Asian
countries is being destroyed
partly through the introduction of
modern cultivars. Conservation of
traditional land races is urgently
needed.
Most wild relatives are
widespread but populations of
several taxa are being lost to
overgrazing in south-west USA
and northern Mexico
Most wild relatives are
widespread but some forms in
Mexico are worthy of
conservation attention.
Breeding relies on a fairly narrow
genetic resource base and efforts
to conserve genetic variability of
the cultivated crop have been
fairly limited.
The species is threatened in the
wild because of logging for its
valuable timber. Commercial
collection of wild nuts is 4
sustainable form of forest
exploitation and is being
promoted in extractive reserves.
2. Uses and Values of Biodiversity
Table 25.1
FAMILY
Musaceae
Myrtaceae
Oleaceae
Palmae
Pedaliaceae
Piperaceae
Rosaceae
Food crops of the world (continued)
SPECIES
Musa acuminata;
M. x paradisiaca
Pimenta dioica
Olea europaea
Cocos nucifera
Phoenix dactylifera
Elaeis guineensis
Sesamum orientale
Piper nigrum
Fragaria x ananassa
Malus pumila
Prunus amygdalus
Prunus armeniaca
Prunus avium
Prunus communis
Prunus domestica
Prunus persica
FOOD
Banana and
Plantain
Pimento
Olive
Coconut
Date
Oil Palm
Sesameseed
Pepper
Strawberry
Apple
Almond
Apricot
Cherry
Pear
Plum
Peach
336
ORIGIN
Wild bananas occur in
SE Asia and the Pacific. The
primary centre for
M. acuminata was the
Malay peninsula.
West Indies and Central
America
Originated as a hybrid in the
eastern Mediterranean
The origin of the coconut is
obscure. Wild types
predominate on the African
and Indian coasts of the
Indian Ocean, and scattered
in Southeast Asia and the
Pacific
A food plant of ancient
cultivation in North Africa
and the Middle East.
West Africa, originally a
species of the transition
zone between savanna and
rain forest.
Possibly Ethiopia or
peninsular India
Wild pepper plants grow in
the Western Ghats of
Malabar, southwestern India
and this is presumed to be
the crop’s centre of origin.
A hybrid between two
American species,
F. chiloensis and F.
virginiana. Both species
were harvested from the
wild and also planted by
Indians before European
settlement. Crossing took
place in Europe in the 18th
century.
Central Asia and Himalayan
region
Central to western Asia
Western China
Western Asia
Central Asia and the
Himalayas
Europe
Western China
CONSERVATION
The genetic base of banana
breeding is narrow. Forest
clearance is threatening the
variability of wild bananas
M. acuminata and other Musa
spp. Protection of wild species in
Asia is an IBPGR conservation
priority.
Olive production is in decline and
the loss of traditionally managed
olive groves has serious
consequences for wildlife in the
Mediterranean region. In Algeria
and Niger the wild olive relative
Olea l/aperrinei is threatened partly
by over-cutting for cattle fodder.
The tendency to plant uniform,
improved hybrids is reducing
genetic variation particularly in
domesticated types.
One wild relative is restricted to
Crete where it is Vulnerable.
In West Africa oil palm groves are
being thinned to make way for
other food crops. Conservation of
the entire genepool in Africa and
parts of Latin America is
considered a priority by IBPGR.
Conservation of wild relatives of
Malus in Europe and Asia is an
IBPGR priority. The Chatkal Mts
Biosphere Reserve, USSR,
conserves apples and various
other fruit trees.
A reserve for the conservation of
almond and other important fruit
trees has been created in the
Kopet Mountains (USSR)
Wild apricots are protected in the
Kopet Mountains Reserve
Protection of wild species in
Europe and Asia is considered a
conservation priority by IBPGR
Table 25.1
FAMILY
Rubiaceae
Rutaceae
Sapotaceae
Solanaceae
Sterculiaceae
Umbelliferae
Vitaceae
Zingiberaceae
Food crops of the world (continued)
SPECIES
Coffea arabica
Citrus aurantiifolia
Citrus grandis
Citrus limon
Citrus x paradisi
Citrus reticulata
Citrus sinensis
Vitellaria paradoxa
Capsicum annuum
Lycopersicon esculentum
Solanum melongena
Solanum tuberosum
Theobroma cacao
Daucus carota
Vitis vinifera
Elettaria cardamomum
FOOD
Coffee
Lime
Pomelo
Lemon
Grapefruit
Tangerine
Orange
Karite nut,
Sheanut
Chili Pepper,
Sweet Pepper
Tomato
Eggplant
Potato
Cocoa
Carrot
Grape
Cardamom
ORIGIN
Ethiopia
Cultivated hybrid with
obscure origins.
Thailand. The origins of
cultivated citrus fruits are
obscure.
Cultivated hybrid with
obscure origins.
The origins of cultivated
citrus fruits are obscure.
The grapefruit is thought to
be a cross between the
pomelo and the sweet
orange.
Southeast Asia
A hybrid, probably
originating in China.
West Africa, grown in
plantations in Ghana and
Nigeria.
Domestication first occurred
in Middle America
The genus is native to
South America. Mexico was
probably the centre of
domestication.
India
The area of domestication is
assumed to be the high
plateau of Bolivia-Peru.
Centre of origin is the
eastern slopes of the Andes
and the centre of cultivation
is Central America.
The species is widespread
in Europe and Asia. The
primary centre of origin for
cultivated forms is thought
to be Afghanistan.
10,000 Old World cultivars
are thought to be derived
from this single wild species
which still occurs in Middle
Asia.
Native to India
Plant Use
CONSERVATION
Coffee grows wild in the
threatened forests of the
Ethiopian massif. Much of the
forest habitat in Ethiopia has been
destroyed. Habitats of wild coffee
are also threatened in Kenya.
Protection of C. arabica in the
wild is a conservation priority.
Protection of wild Citrus species
in Asia is a conservation priority.
Protection of wild Citrus species
in Asia is a conservation priority.
Protection of wild Citrus species
in Asia is a conservation priority.
Protection of wild Citrus species
in Asia is a conservation priority.
Wild peppers are still collected
and sold locally. A large number
of yet unexploited varieties exist
in the Tropics. More collection for
seed banks is needed.
The wild relatives of the tomato
have limited ranges. The crop’s
wild gene pools are prone to
erosion by habitat destruction.
There are over 150 wild species
of potato, many of which have
limited natural distributions;
3,000-5,000 varieties are
recognised by farmers in the
Andes. Conservation of
genetically valuable local varieties
is being carried out at the
International Potato Centre in
Peru.
Cultivated varieties suffer from a
lack of genetic variation. Forests
harbouring genetic diversity in the
wild are being rapidly destroyed.
Wild relatives are suffering
genetic erosion in the USA.
Collection from the wild
contributes to the commercial
trade.
Source: Compiled from multiple sources. Species list based on Prescott-Allen, C. and Prescott-Allen, R. 1990. How many plants feed the world?
Conservation Biology 4(4):365-374. Historic, cultivation and nomenclatural data from Simmonds, N.W. (Ed.) 1976. Evolution of Crop Plants.
Longman Scientific and Technical. Conservation data from multiple sources.
337
2. Uses and Values of Biodiversity
Most of the world’s major food crops were domesticated
and widely dispersed by 2,000 years ago. Diversification
continued during colonial periods from the spread of Roman
civilisation through to European settlement of the tropics.
Colonial expansion undoubtedly contributed to the loss of
genetic diversity of cultivated plants in the New World as
a result of devastation of farming communities by invasion
and disease. It has been suggested, however, that more
varieties arose as a result of crop interchange between
continents and islands during colonial expansion than were
lost through cultural disintegration (Plucknett et al., 1987).
Maize and cassava were, for example, introduced to Africa
by the Portuguese in the 16th century and diversified as
they were grown under a new range of ecological and
cultural conditions.
The history of food crops is complex and the exact origins
of some cultivated food plants are obscure. Nevertheless,
the geographical origins of major crops can be traced back
through time.
From the earliest stages of agriculture, regions of diversity
developed which remain important centres of crop
biodiversity today. In addition, there are a number of minor
centres of origin where a few crops can be related to their
initial domestication in particular localities.
The Russian botanist N.I. Vavilov first described and
mapped centres of diversity for individual crops which he
believed represented their centres of origin. It became
apparent that the centres of diversity of different crops
coincided to give remarkable concentrations of crop plant
variation. Vavilov (1951) ultimately recognised eight such
centres; later authors have modified the centres and
identified new ones (Fig. 25.1).
In general the concept of ‘Vavilov Centres’ where a centre
of current diversity is taken to indicate the centre of origin
of crops is now considered an _ oversimplification.
Nevertheless, geographical concentrations of crop variation
are real and these areas are of immense conservation
importance. The reasons for the diversity are: the great age
of cultivation in such centres, the wide range of ecological
conditions and farming practices found within them, and the
processes of natural selection caused by the presence of
many different pathotypes of pests and diseases and by the
variable ecological conditions. Some features of one of
Vavilov’s Centres of Diversity, the Ethiopian centre, are
described in the case study below.
Crop genetic resources
Genetic resources can be defined as the genetically
transmitted characteristics of organisms which are of actual
or potential value to people. Such characteristics may
include rapid growth, high yields, disease- and pest-
resistance and environmental adaptation. The genetic
resources of crop plants represent the total genetic diversity
of cultivated species and their wild relatives, much of which
is of immense value in crop breeding programmes. Many
of the species from which crop plants have been selected
continue to survive in the wild today. These, together with
closely related species, comprise the wild relatives of crops.
They continue to evolve under natural conditions and
338
provide a largely untapped reservoir of genetic diversity.
Gene flow between cultivated crops and wild relatives
continues to occur today, and is encouraged in areas where
traditional forms of agriculture are still practised. In
Mexico, for example, some traditional farmers still utilise
teosinte, the closest wild or weedy relative of maize, to
increase corn yields. The weedy plants are allowed to
remain within or near cultivated maize populations so that
natural crosses may occur and produce fertile hybrid stock
that can be selected for desirable characteristics.
The genetic resources of wild crop relatives can be
classified according to the ease with which the species can
be crossed with the cultivated crop. The primary gene pool
consists of relatives that are interfertile and hybridise
readily with the cultivated crop. Wild forms of cocoa
Theobroma cacao occurring in the Amazon forests can, for
example, be crossed readily with cultivated cocoa and
constitute the ‘primary gene pool’ for the crop. The same
applies to certain wild relatives of maize Zea mays.
The ‘secondary gene pool’ consists of species which can be
crossed using conventional breeding methods but crossing
is difficult and only a small proportion of first generation
progeny may be fertile. The secondary gene pool of maize
includes, for example, wild relatives Zea perennis and
Tripsacum species. The ‘tertiary gene pool’ of a crop plant
consists of species that are more distantly related. The
genetic diversity in tertiary gene pools can only be utilised
by experimental techniques in plant breeding such as using
another species as a bridge.
The most common use of wild genetic resources in crop
breeding programmes has been in the introduction of
resistance to pests and diseases. Wild tomato species,
Lycopersicon pimpinellifolium and L. peruvianum have, for
example, been used in breeding programmes to confer
resistance to various forms of bacterial wilt. Genes from
wild relatives of the tomato have also conferred resistance
to a range of viruses, moulds, and other pests. Likewise,
wild potato relatives have been crossed with cultivars for
about a hundred years, the wild species yielding genes for
resistance to viruses, bacterial wilt, nematodes, aphids and
a range of other potato disorders.
In addition to wild crop relatives a second important
storehouse of genetic crop diversity is the range of variation
shown by ‘land races’. These are races or populations of
crops that have become adapted under natural and artificial
selection processes to the local conditions under which they
are cultivated. Land races have not been deliberately bred
but have been developed over centuries of traditional
agriculture. They are now being explored as a source of
genetic material for crop improvement programmes. Recent
work in the Himalayan foothills of north-east India has, for
example, revealed a large number of primitive rice cultivars
with resistance to major pests and diseases including
bacterial blight, tungro virus, gall midge and stem borer.
Genetic erosion
The evolution of food crops under centuries of
domestication has increased variation as seen in the main
regional centres of crop diversity. But the development of
Plant Use
Figure 25. 1 Regions of diversity of crop plants
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339
2. Uses and Values of Biodiversity
high-yielding modern cultivars for intensive agriculture is
now rapidly reversing this trend, leading to a dangerous
reliance on genetically uniform crops.
Genetic erosion, or the loss of genetic diversity, of the
world’s food plants is an issue of serious concern with
implications for the long-term maintenance of global food
supplies. At a time when more genetic diversity is needed
in crop breeding programmes to increase food production
this diversity is rapidly disappearing or has already been
lost. Economic aspects of this process are discussed in
Chapter 27.
Various factors contribute to genetic erosion. The
worldwide threats to wild species through habitat
destruction and modification have an obvious impact on
wild crop relatives. Habitat destruction is having a direct
effect on, for example, wild forms of cocoa: large parts of
the centre of genetic diversity of Theobroma cacao in
Colombia, Ecuador and Peru have been destroyed as a
result of petroleum exploration and exploitation, and by
agricultural expansion. Similarly, around 90% of the
Ethiopian highland forests, which harbour wild coffee
Coffea arabica, have been destroyed.
Of equal concern is the loss of old land races through
replacement by modern highly-bred crop cultivars. This
may lead to improvements in yield but also results in
increased reliance on agrochemicals and all the problems
associated with monoculture cultivation.
The extent of genetic erosion differs for various crops. In
general, the wild relatives of cereals are widespread, weedy
and thrive in disturbed ground. There is some evidence of
genetic erosion of wild relatives, however, and conservation
attention is a priority for those of rice, wheat and maize.
The loss of local land races for these major cereals has been
a particularly serious problem in various parts of the world.
The introduction of new high-yielding varieties of wheat
has, for example, caused severe genetic erosion in Turkey,
Iraq, Afghanistan, Pakistan and India. In Greece, 95% of
the native varieties of wheat have been lost in 40 years
(Davies, 1991).
The wild relatives of root crops are also suffering loss of
genetic diversity. More than half the wild species in the
genera Solanum (potato) and Manihot (cassava) are
narrowly endemic in South, Central and Middle America
(FAO, 1984). Conservation of centres of potato diversity is
an urgent concern, as some species have already become
extinct. Diversity of natural populations of Manihot species
is declining owing to conversion of their habitats to pasture
and elimination of the plants, which are poisonous to
grazing animals.
All the wild relatives of the tomato Lycopersicon esculentum
have limited natural distributions. Clearance of habitats for
agriculture, housing and industry has led to the loss of wild
populations of the tomato species Lycopersicon hirsutum
and L. peruvianum. As yet no in situ conservation areas
have been established for wild tomato plants. Loss of
genetic resources from the wild can also be seen with
Brassica oleracea. This species, native to southern England,
western France and northern Spain, provides a number of
340
cultivars including cabbages, cauliflowers and Brussels
sprouts. The populations related to B. oleracea form a
group of about 12 perennial species (often considered to be
subspecies of B. oleracea) most of which are endemic to
the western Mediterranean. These species are composed of
small populations often reduced to a few specimens and
isolated geographically (Valdes, 1991). Several of the
species are considered by IUCN to be Rare, Endangered or
threatened. Other wild taxa in the group may not be
endangered as species although many of their populations
are. In many cases, wild Brassica populations in the
Mediterranean are protected by the inaccessible nature of
their rocky habitats but elsewhere they are threatened by
competition from maquis or garrigue scrub or by habitat
destruction (Olivier, 1991).
Examples of genetic erosion in wild populations and
traditional cultivars can be found in all groups of food crops
throughout the world. Summary information is given in
Table 25.1. Lack of knowledge of intra-specific genetic
variation remains a problem in detecting the degree of
threat to plant gene pools, but certain priorities have been
established for international conservation action. The
conservation of plant genetic resources is discussed in Part
3:
Case study: the Ethiopian centre of crop genetic
diversity
Ethiopia represents one of the world’s eight major centres
of crop plant diversity. It is the probable area of
domestication for many crops and for others, where no wild
relatives are known within the country, it is a secondary
centre of diversity. The rich variation within crop plants
results from the highly dissected topography of the country
allowing crops to evolve in isolation under primitive
agricultural conditions, together with the ancient and very
diverse cultural history of the country. The geographical
position of Ethiopia, at the crossroads between the Near
East and Indian centres of diversity, also accounts for the
genetic richness of the country’s crop plants.
Below are described 12 widespread crops which are
believed to have their centres of diversity within the region
along with three other Ethiopian crops - chat, ensete and
noog - which have originated and evolved within the
country (Engels et al., 1991).
In addition to the cultivated crop species, there are many
wild plants used for food in Ethiopia, particularly in times
of food shortage. Some have considerable potential as new
crop plants. One such example is the yeheb nut Cordeauxia
edulis. This species is endemic to eastern Ethiopia and part
of Somalia. It has long been valued for its highly nutritious
nuts but is now Endangered in the wild because of over-
exploitation and overgrazing.
Many other plant species are of local importance. Medicinal
plants, for example, remain important sources of drugs for
nearly 80% of the Ethiopian population. Most of Ethiopia’s
major medicinal plants are not in cultivation.
Plant species have a wide range of other uses in Ethiopia.
Fuelwood, for example, provides for over 90% of the
country’s total energy consumption. Timber, resins, gums,
cosmetics, perfumes, dyes, inks, fibres, and forage are all
derived from wild plants. Ethiopia’s rich plant biodiversity
is clearly of immense importance. The country’s flora and
vegetation types are, however, still incompletely known and
information on the conservation status of individual species
is sparse. The almost complete deforestation of Ethiopian
highlands, changes in land-use and agricultural practices are
undoubtedly having serious consequences for both wild and
cultivated plant diversity.
Concern about the loss of plant genetic resources led to the
establishment of the Plant Genetic Resources Centre
(PGRC/E) in 1976. The Centre is involved in exploration,
collection and preservation of crop germplasm, together
with the provision and exchange of germplasm for crop
breeding programmes. Collection of genetic resources
includes the collection of land races from drought-prone
areas for storage at seed reserve centres and redistribution
to farmers when required. This is an insurance measure to
prevent major losses of crop genetic diversity by
consumption of seed in times of famine or the replacement
of traditional varieties by imported seeds distributed through
relief agencies. The involvement of farmers in the
conservation of germplasm is being considered. Already
measures are under way to conserve semi-cultivated coffee
on peasant farms and ‘backyard’ coffee in cooperatives, as
part of the national coffee conservation programme.
Coffee (Coffea arabica; Rubiaceae) Coffea arabica
accounts for over 80% of the world’s coffee production.
Almost the entire diversity of this crop originated in
Ethiopia, mainly in the south-western rain forest area.
C. arabica still occurs as a wild plant in these moist
montane forests and as a semi-wild or cultivated crop in the
same areas. In drier parts of Ethiopia it is grown as an
irrigated crop and elsewhere as a garden plant often mixed
with fruit trees and herbs. There is extremely high genetic
diversity within Ethiopian coffee but this rich diversity is
under considerable threat. Deforestation, replacement of
primitive coffee populations by other crops and changing
patterns of land-use are leading to severe problems of
genetic erosion.
Barley (Hordeum vulgare; Gramineae) Ethiopia’s third
most important cereal crop, barley, was introduced from the
Near East in ancient times. Ethiopia is a secondary centre
of diversity for the species and the crop has developed
Many important and unique characteristics within the
country. Extensively grown land races can still be found,
but genetic erosion is resulting from replacement by other
cereals.
Sorghum (Sorghum bicolor; Gramineae) Ethiopia and
Sudan are assumed to be the primary centre of origin and
diversity of sorghum. This cereal is probably the most
diverse of all Ethiopia’s crops. It is grown in a wide range
of ecological conditions throughout the country. Disease-,
pest- and drought-resistance have all been reported. Use of
improved local land races and imported varieties together
with replacement by maize and other crops are causing the
loss of genetic diversity.
Wheat (Triticum spp.; Gramineae) Ethiopia represents a
secondary centre of diversity for wheat. Durum wheat is the
341
Plant Use
main type grown. It exhibits high phenotypic diversity
within the country and agronomically important genes have
been located in Ethiopian germplasm. Genetic erosion is
occurring because of replacement by other crops.
Teff (Eragrostis tef,; Gramineae) Teff is the most widely
grown crop in Ethiopia. It is used mainly for making a
pancake-like bread called ‘injera’ and also to make porridge
and alcoholic drinks. The straw is used as a cattle feed and
in house construction. Elsewhere teff is cultivated only in
North and South Yemen although E. tef has a wide
distribution in Africa. Ethiopia is the centre of origin for
teff and domestication is thought to have first taken place in
the northern highlands. Over 50 Eragrostis spp. occur in
Ethiopia, of which 14 are endemic. At least 35 land races
of teff are known in the country. Genetic erosion is not a
problem for the crop which is still expanding its acreage.
Improved varieties are not being introduced.
Niger seed, noog (Guizotia abyssinica; Compositae) Noog
is the most important oil crop in Ethiopia, and the area
under cultivation is expanding. It is thought that the crop
originated in the highlands of Ethiopia and that it was one
of the earliest crops to be domesticated within the country.
It probably originated from the wild species Guizotia
scabra, which now frequently occurs as a weed of noog
fields. Genetic erosion is not currently a problem faced by
noog, but improvement of agricultural practices may lead to
genetic erosion of G. scabra.
Linseed (Linum usitatissimum; Limaceae) Linseed is the
second most important oil crop in Ethiopia. It was
introduced in ancient times from Asia, and Vavilov
considered Ethiopia to be a centre of flax diversity. This
diversity is seriously threatened by genetic erosion.
Sesame (Sesamum indicum; Rubiceae) Ethiopia is likely
to be the centre of origin of sesame, but this remains
uncertain. Economically, sesame is the third most important
oil crop in Ethiopia. The crop exhibits considerable
diversity but is facing critical genetic erosion.
Castor bean (Ricinus communis; Euphorbiaceae) Castor
bean is not cultivated as a commercial crop in Ethiopia. It
is widely distributed throughout the country as a wild plant
or weed and is used as a medicinal plant or source of oil
for lighting. Phenotypic diversity is enormous, and this has
led to suggestions that the cultivated castor bean might be
of Ethiopian origin. There is no threat of genetic erosion at
present.
Pea (Pisum sativum; Leguminosae) A unique subspecies
occurs in Ethiopia. It has been suggested that Ethiopia is
one of four possible centres of diversity of the pea.
Phenotypic diversity is rather limited and the degree of
genetic erosion is expected to be low.
Chickpea (Cicer arietinum; Leguminosae) Chickpea is an
ancient crop in Ethiopia, and the country is a centre of
diversity for the cultivated plant. Phenotypic variation is
considerable, and initial testing has shown some disease
resistance and drought tolerance. Genetic erosion is not a
significant threat.
2. Uses and Values of Biodiversity
Lentil (Lens culinaris; Leguminosae) The lentil was an
early introduction into Ethiopia from west Asia. The crop
shows a high degree of diversity. Genetic erosion is
expected as the acreage of lentils is declining.
Ensete (Ensete ventricosum; Musaceae) Ensete is a crop
species unique to Ethiopia. Both wild and cultivated forms
occur throughout the country wherever there is sufficient
moisture. The pseudocorm is processed to form a staple
food; other parts of the plant are used as fodder, fuel,
packing material, to wrap bread during cooking and to
make ropes. The crop shows considerable variation and
over 70 named varieties have been described. Bacterial wilt
and drought are contributing to genetic erosion.
Chat (Catha edulis; Celastraceae) The leaves of this
evergreen shrub are used as a stimulant. The plant was first
domesticated in Ethiopia. Cultivation is now expanding and
is leading to the replacement of coffee in the eastern part of
the country. No genetic erosion is currently taking place.
TIMBER
Wood is one of the basic commodities utilised worldwide
that is still predominantly harvested from the wild. It
provides the primary source of fuel in many developing
nations, shelter in traditional home-building and
sophisticated construction, and the basis for the international
pulp and paper industry. Wood is one of the most important
commodities in international trade and accounts for a
particularly significant proportion of the export earnings of
developing tropical countries. In 1989 the total worldwide
value of wood exports was around US$6 billion.
Table 25.2 provides figures for wood production and trade
for 1989. The bulk of the wood in world trade comes from
temperate sources, with the major exporters being USA, the
former USSR, and Canada for logs and sawnwood and
USA, the former USSR and Finland for plywood. The main
tropical source countries are Malaysia, Papua New Guinea
and Gabon for logs, and Malaysia and Indonesia for
sawnwood and plywood.
Within developing countries there is a trend towards value-
added processing in the timber industry. The export of
timber in log form is increasingly being restricted partly to
retain wood within the country for further processing and
partly as a conservation measure. Nevertheless, logs still
account for a significant proportion of world trade.
Overall, developing countries still retain a relatively small
proportion of the financial value of their timber resources
despite increasing industrialisation of the forestry sector.
The trade imbalance remains heavily in favour of the
developed nations. It has been shown, for example, in a
recent study that 65-90% of the growth in value of tropical
forest products occurs in consumer countries, made up of
operating costs, tax revenue and profits (Oxford Forestry
Institute, 1991).
In general it is difficult to assess the extent to which timber
for domestic consumption or international trade is derived
from plantations. Industrial timber plantations mainly
consist of conifers which lend themselves well to cultivation
342
as pure crops. Relatively few hardwoods have been
cultivated as plantation timbers. The majority of hardwoods
in international trade are derived from natural forests which
are subject to varying degrees of management. In tropical
regions relatively few examples of successful forest
management for sustainable timber production are known.
Detailed information on levels of production and trade in
individual timber species is scarcely assembled at an
international level. The conventional division of timber
products into hardwoods (non-coniferous) and softwoods
(coniferous), for trade purposes, disguises the great
diversity of wood as a natural product. Timber species
richness is particularly high in tropical regions. Ghana, for
example, has 674 tree species reaching timber size and
timber from about 60 of these has been exported in the past
20 years. Peninsular Malaysia has at least 3,000 tree
species of which over 400 have been traded on international
markets. Developing countries are attempting to diversify
their timber exports by promoting lesser known species but
consumer demand remains conservative in importing
countries. Potentially valuable timber resources are under
threat in many parts of the world through inadequate
management, habitat loss and over-harvest (Table 25.3).
Ghana Timber Species Case Study
Timber is Ghana’s third most important export commodity
after cocoa and minerals. Ghana’s share of the world’s
tropical timber trade is about 1% and it accounts for about
3% of West European imports of tropical hardwoods.
Europe takes over 90% of Ghana’s timber exports. Logs
are predominantly exported to Germany and the UK;
Germany and Ireland are leading sawn timber importers and
sliced veneer goes mainly to Germany.
The timber export trade began a century ago, concentrating
on species of Entandrophragma and Khaya, the so-called
African Mahoganies. In total around 674 tree species reach
timber size in Ghanaian forests and timber of about 60 of
these has been exported over the past 20 years. Commercial
exploitation over the past century, together with the
reduction of natural forest from 8 million ha to below 2
million ha, has placed considerable pressure on the
commercial timber species.
The report of the Fifth Session of the FAO Panel of Experts
on Forest Gene Resources drew attention to the fact that in
Ghana, “some of the most valuable commercial species
Pericopsis elata, Gossweilerodendron balsamiferum, Lovoa
trichilioides, Entandrophragma utile, Nauclea diderrichii,
Terminalia ivorensis, T. superba, Antiaris africana,
Triplochiton scleroxylon and Hallea ledermannii, are
threatened with extinction in their areas of natural
distribution because of massive exploitation."
More recently a full inventory of Ghana’s timber resources
has been carried out in a project funded by the UK’s
Overseas Development Administration. Information from
this study suggests that immediate, serious problems of
over-logging apply to timbers of the Meliaceae, especially
Khaya ivorensis, and also to Pericopsis elata. Taxa such as
Terminalia and Triplochiton scleroxylon are, in fact,
regenerating well in disturbed forest and are relatively fast
Table 25.2 Wood production and trade, 1989
Plant Use
ROUNDWOOD PRODUCTION! TIMBER PRODUCTION PAPER + NET
FUEL AND INDUSTRIAL SAWNWOOD WOOD-BASED PAPERWOOD TRADE2
TOTAL CHARCOAL ROUNDWOOD & SLEEPERS PANELS PRODUCTION ROUNDWOOD
m3 m3 m3 m3 m3 metric tons m3
ASIA 1.07E+09 793913140 273434972 106701500 27453931 5122400 110238000
Afghanistan 6104000 4609000 1495000 400000 1400
Bahrain 36000000
Bangladesh 30144992 29272000 873000 79000 8000 96000
Bhutan 3224000 2946000 278000 5000 —7000
Brunei 294000 79000 215000 90000
China 274589952 177610016 96980000 24958000 3650000 15336000 13382679000
Cyprus 78300 22300 56000 57300 22000
Hong Kong 187000 187000 248000 12000 40000 699637000
India 269450752 245126992 24324000 17460000 441700 1940000 902324000
Indonesia 175730496 136079008 39651488 10390500 8838427 974000 —1131000
Iran, Islamic Rep 6829000 2453000 4376000 163000 54200 78000 117300000
lraq 149000 99000 50000 8000 3000 28000 1200000
Israel 118000 11000 107000 148000 180000 212500000
Japan 31935904 571000 31364896 30542000 8993000 26809000 51809890000
Jordan 9000 5000 4000 10000 16900000
Kampuchea, Dem 5803000 5236000 567000 43000 2300
Korea, Dem People’s Rep 4761000 4161000 600000 280000 80000 70600000
Korea, Rep 6803000 4491000 2312000 4014000 1453000 4018000 7384300000
Kuwait 60493000
Laos 3972000 3660000 312000 16000 10000 —34000
Lebanon 503000 482000 21000 27000 46300 37000 15500000
Malaysia 50536688 8258000 42278688 8275000 1630000 70000 282530000
Mongolia 2390000 1350000 1040000 470000 3500
Myanmar 22287008 17407008 4880000 463700 15200 8000 —360000
Nepal 17804000 17244000 560000 220000 2000
Oman 22500000
Pakistan 24408992 23226000 1183000 751000 93500 151000 34700000
Philippines 38503008 33075008 5428000 950000 425000 334000 397895000
Qatar 35700000
Saudi Arabia 221300000
Singapore 206000 489000 10000 170688000
Sri Lanka 8988400 8302000 686400 20000 9604 28000 —33000
Syrian Arab Rep 48300 14800 33500 9000 26700 19000 22396000
Thailand 38734000 34115008 4619000 1279000 257000 520000 1410029000
Turkey 15449000 9721000 5728000 4923000 781000 400000 467570000
Viet Nam 27188000 23776000 3412000 354000 40100 56000 41100000
Yemen, People’s Dem Rep 324000 324000 5199000
USSR* 382099968 80700000 301399808 100000000 14635000 10654000 136949000
EUROPE 368250792 53890800 314360008 86292500 38232200 66749000 = —29257000
Albania 2330000 1608000 722000 200000 12000 24000
Austria 16086000 1413000 14673000 7054000 1538000 2754000 4830078000
Belgium and Luxembourg 4757000 572000 4185000 1114000 2247000 1237000 4127038000
Bulgaria 4455000 1810000 2645000 1283000 496000 438000 134661000
Czechoslovakia 18552000 1532000 17020000 4993000 1441000 1312000 50552000
Denmark 2118000 467000 1651000 861000 331200 326000 382296000
Finland 46262000 2984000 43278000 7763000 1482000 8752000 6783253000
France 43726992 10436000 33290992 10559000 3018000 6754000 1756597000
German Dem Rep 10897000 626000 10271000 2555000 1179000 1351000 539809000
Germany, Fed Rep 35332000 3656000 31676000 11405000 8529000 11259000 _ 3903311000
Greece 3289000 2320000 969000 355200 398000 282000 244697000
Hungary 6609000 2949000 3660000 1256500 449000 504000 1215830000
Iceland 1200000
Ireland 1527000 50000 1477000 300000 236000 34000 7688000
Italy 8846000 4177000 4669000 1998000 4342000 5555000 6458978000
Malta 400000
Netherlands 1331000 116000 1215000 465000 94000 2570000 1327486000
Norway 11039000 936000 10103000 2491800 611000 1789000 1667565000
Poland 22348992 3123000 19226000 4963000 1845000 1406000 247260000
Portugal 10341800 597800 9744000 1650000 1025000 740000 819269000
Romania 19306992 2790000 16517000 2851000 1525000 819000 56884000
Spain 17182000 2384000 14798000 2724000 2295000 3446000 1940367000
Sweden 55704016 4424000 51280016 11487000 1307000 8362000 8207344000
Switzerland 4562000 850000 3712000 1282000 918000 1259000 961168000
United Kingdom 6462000 209000 6253000 2191000 1676000 4475000 901083000
Yugoslavia 15186000 3861000 11325000 4491000 1238000 1302000 1072728000
NORTH & CENTRAL AMERICA 771385180 172135000 599250156 166232942 40996200 89698000 . —33376000
Bahamas 115000 115000 1400 300000
Barbados 500000
Belize 187600 126000 61600 14300 —8000
Canada 176976016 6834000 170142016 59224992 6913000 16555000 4263107000
Costa Rica 4037000 2886000 1151000 515400 57500 18000 —4000
Cuba 3122000 2511000 611000 130100 149000 168000 5700000
Dominica 700000
Dominican Rep 982300 976000 6300 0 10000 40500000
El Salvador 4440000 4320000 120000 54000 17000 100000
Guadeloupe 17000 15000 2000 1000 900000
Guatemala 7604000 7490000 114000 83000 6000 17000 —12000
Haiti 5727000 5488000 239000 13800
Honduras 6056000 5172000 884000 441000 10000 —21000
Jamaica 218000 13000 205000 40000 0 4000 200000
Martinique 11900 10000 1900 950
Mexico 22628992 15204000 7425000 2410000 645300 3375000 19194000
Netherlands Antilles 300000
Nicaragua 3972000 3092000 880000 222000 3400 200000
Panama 2047300 1708000 339300 18000 12000 20000 1700000
Trinidad and Tobago 75200 22000 53200 23000 1100000
United States 533167872 116268000 416899840 103040000 33200000 69514000 3442768000
343
2. Uses and Values of Biodiversity
Table 25.2 Wood production and trade, 1989 (continued)
ROUNDWOOD PRODUCTION TIMBER PRODUCTION PAPER + NET
FUEL AND INDUSTRIAL SAWNWOOD WOOD-BASED PAPERWOOD TRADE
TOTAL CHARCOAL ROUNDWOOD 4& SLEEPERS PANELS PRODUCTION ROUNDWOOD
m3 m3 m3 m3 m3 metric tons m3
SOUTH AMERICA 335574900 234266700 101308208 26624292 4112500 7572000 4763000
Argentina 10819000 4332000 6487000 1446000 354000 917000 3299000
Bolivia 1556500 1301000 255500 95000 3900 2000
Brazil 255455008 182806000 72649008 18178992 2892000 4806000 25854000
Chile 16864000 6540000 10324000 2713000 282000 445000 —4679000
Colombia 18478992 15806000 2673000 721200 113000 501000
Ecuador 9728000 6642100 3085900 1491600 145400 35000
French Guiana 253600 65600 188000 19000 te) —5000
Guyana 228000 19000 209000 57000 0 —22000
Paraguay 8394000 5288000 3106000 906000 106500 11000
Peru 8785800 7669000 1116800 541500 36000 260000 100000
Suriname 235000 20000 215000 73000 9700 —11000
Uruguay 3295000 3038000 257000 57000 10000 70000 400000
Venezuela 1482000 740000 742000 325000 160000 524000 11800000
OCEANIA 39795692 8738000 31057692 6004573 1774100 2605000 11923000
Australia 20040992 2886000 17154992 3612000 1081000 1870000 —5814000
Fiji 306800 37000 269800 93600 16100 —148000
French Polynesia 1300000
New Caledonia 12100 12100 5400 400000
New Zealand 10557000 50000 10507000 2131000 658000 735000 2830000
Papua New Guinea 8231000 5533000 2698000 117000 19000
Solomon Islands 449000 138000 311000 16000 0
Tonga 4600 4600 1573
Vanuatu 63200 24000 39200 7000
Western Samoa 131000 70000 61000 21000 0
AFRICA 498508268 441864308 56644000 8829600 1904423 2460000 —5292000
Algeria 2131000 1874000 257000 12800 49500 120000 210000000
Angola 5402000 4335000 1067000 5000 2000 15000
Benin 5000000 4738000 262000 11000
Botswana 1321000 1239000 82000
Burkina Faso 8526400 8141000 385400 750
Burundi 4083000 4034000 49000 3000
Cameroon 12850000 10142000 2708000 653000 80000 5000
Central African Rep 3455000 3055000 400000 52000 3900
Chad 3936000 3380000 556000 1000
Congo 3300000 1776000 1524000 46000 54400
Céte d'Ivoire 13243000 9830000 3413000 775000 266000 —550000
Djibouti () te) ()
Egypt 2266000 2161000 105000 80000 160000 200100000
Equatorial Guinea 607000 447000 160000 51000 10000
Ethiopia 39640000 37884000 1756000 34000 15300 10000
Gabon 3700000 2478000 1222000 126000 228000 —913000
Gambia, The 921600 901000 20600 1000
Ghana 17168992 16068000 1101000 537000 53000 —201000
Guinea 4669000 4022000 647000 90000 0 —8000
Guinea-Bissau 567000 422000 145000 15700
Kenya 35650000 33884000 1766000 185000 52200 108000
Lesotho 596000 596000 32700000
Liberia 5960000 4800000 1160000 411000 5000 —701000
Libya 642000 536000 106000 31000 6000 29800000
Madagascar 7856000 7049000 807000 234000 5000 6000 —2000
Malawi 7621000 7275000 346000 31000 6200
Mali 5515800 5163000 352800 12800
Mauritania 12000 7000 5000
Mauritius 31000 16500 14500 4600 0 100000
Morocco 2110600 1363600 747000 83000 147000 109000 600500000
Mozambique, People’s Rep 16027000 15022000 1005000 35750 2800 2000 —1000
Niger 4418000 4146000 272000
Nigeria 108298000 100430000 7868000 2712000 233000 73000 984000
Réunion 33300 31000 2300 2200 1600000
Rwanda 5842000 5602000 240000 13000 1723
Sao Tome and Principe 9000 9000 5400
Senegal 4391000 3786000 605000 11000 25300000
Sierra Leone 3014000 2874000 140000 12000
Somalia 6986000 6896000 90000 14000 0 300000
South Africa 19360992 7078000 12283000 1873000 398000 1636000 26257000
Sudan 22198992 20112000 2087000 12500 1500 10000
Swaziland 2223400 560000 1663400 136000 8000 71830000
Tanzania 33102992 31114000 1989000 156000 14700 28000
Togo 866000 683000 183000 5000 200000
Tunisia 3177500 3015000 162500 20000 97000 82000 15800000
Uganda 14365000 12507000 1858000 28100 3300 2000
Zaire 35348000 32557008 2791000 121000 52500 2000 —117000
Zambia 12204000 11565000 639000 76000 8200 4000
Zimbabwe 7861700 6269200 1592500 190000 26200 82000 —2000
Source: FAO Yearbook 1989, Forest products.
Note: ' Roundwood refers to all wood in the rough destined for either fuel or industrial uses. It includes sawlogs, veneer logs and pulpwood. 2 Net
trade is the balance of imports minus exports. * Former USSR.
growing. A quantitative measure of the status of various the resource life of Pericopsis elata is already zero.
timbers was provided by an estimation of their resource life
undertaken as part of the inventory project. Estimates based The various assessments of frequency, resource life and
on the results of the project for resource data combined conservation status of a number of Ghana’s commercial
with information on growth and extraction rates suggest that timbers are summarised in Table 25.4. This table also
344
Table 25.3
FAMILY
Anacardiaceae
Apocynaceae
Aquifoliaceae
Araliaceae
Araucariaceae
Betulaceae
Bignoniaceae
Bombacaceae
Boraginaceae
Cercidiphyllaceae
Chenopodiaceae
Compositae
Cupressaceae
SPECIES
Astronium
urundeuva
Schinopsis
brasiliensis
Aspidosperma
polyneuron
Ilex paraguaiensis
Didymopanax
morototoni
Araucaria
angustifolia
Araucaria
cunninghamii
Araucaria hunsteinii
Alnus acuminata
Tabebuia
impetiginosa
Zeyhera tuberculosa
Bombacopsis
quinata
Cordia milleni
Cercidiphyllum
japonicum
Atriplex repanda
Brachylaena
huillensis
Cupressus atlantica
Cupressus
dupreziana
Juniperus
bermudiana
Juniperus procera
DISTRIBUTION
Brazil, Argentina,
Paraguay
North-eastern Brazil
Brazil, Argentina,
Paraguay and Peru
South America
Central and South
America and
Caribbean islands
Brazil and Argentina
Irian Jaya and Papua
New Guinea
Papua New Guinea
Mexico, Central
America and the
Andes
Brazil
South-eastern Brazil
Tropical America
Tropical Africa
Japan and China
Chile
Central Africa
Morocco
Algeria
Bermuda
Arabia and Tropical
Africa
345
Endangered tree species and provenances
STATUS
Endangered
Suffering a slow
decline
Over-exploited
Abundant but in need
of conservation
attention
Endangered in parts
of its range
Endangered in parts
of its range
Endangered in parts
of its range
In danger of genetic
impoverishment
Suffering a slow
decline
Threatened
Severely threatened
at the provenance
level
Rare in Kenya
Provenances are
endangered in China
Endangered in parts
of its range
Endangered
Endangered
Endangered
Outlying populations
endangered
Plant Use
THREATS
Exploitation for
timber, tannin and
medicinal purposes
Intensive exploitation
and habitat
conversion
Leaves used to
produce mate, a tonic
and stimulant drink
Heavy utilisation
Excessive exploitation
of wild stands
Habitat destruction,
logging, low natural
regeneration
Shifting agriculture,
fire, over-exploitation
Substitution with
introduced fast-
growing species
Agriculture, livestock
and charcoal
production
Excessive felling and
forest clearance
Forest clearance and
utilisation of species
for timber and
medicinal purposes
Over-utilisation for
livestock
Habitat clearance and
excessive felling
Exploitation and
increasing human
pressure
Grazing and
exploitation for
firewood
Approx. 90% of the
trees died between
1944 and 1950 as a
result of severe insect
infestation
Fire, browsing
pressure particularly
from buffalo and
elephant, logging and
plantation
development
2. Uses and Values of Biodiversity
Table 25.3
FAMILY
Ebenaceae
Euphorbiaceae
Fagaceae
Hamamelidaceae
Irvingiaceae
Lauraceae
Lecythidaceae
Leguminosae
SPECIES
Diospyros hemiteles
Joannesia principes
Fagus longipetiolata
Liquidambar
styraciflua
Irvingia gabonensis
Aniba duckei
Ocotea porosa
Bertholetia excelsa
Acacia albida
Acacia caven
Acacia tortilis ssp.
raddiana
Acacia tortilis ssp.
tortilis
Anadenanthera
macrocarpa
Caesalpinia dalei
Dalbergia nigra
Dipterix alata
Gigasiphon
macrosiphon
Gossweilero-
dendron
balsamiferum
Machaerium
villosum
DISTRIBUTION
Mauritius
Brazil
China
Southern USA and
Central America
Tropical Africa
Amazon region
Brazil
Bolivia, Brazil,
Colombia, Peru and
Venezuela
Sub-Saharan Africa
and parts of the
Middle East
Bolivia, Argentina,
Uruguay and Chile
North Africa, Egypt,
Israel, Jordan, Saudi
Arabia
Somalia, Ethiopia,
Sudan, Egypt, Israel,
Arabia
Brazil, Argentina,
Peru, Bolivia,
Paraguay
Kenya
Brazil
Brazil
Kenya and Tanzania
Nigeria to Zaire
Brazil
346
Endangered tree species and provenances (continued)
STATUS
Endangered; one
individual remains in
the wild
Threatened
Endangered in parts
of its range
Endangered in parts
of its range
In danger of genetic
impoverishment
Threatened
Some stands
threatened in Israel
Endangered in parts
of its range
Endangered in parts
of its range
Endangered in parts
of its range
Suffering a slow
decline
Endangered
Endangered
Threatened
Rare or Endangered
Likely to be
endangered in parts of
its geographic range
Threatened
THREATS
Cultivation, illegal
wood cutting, monkey
and deer damage,
invasive plant species
Forest clearance and
commercial
exploitation
Land clearance for
agriculture and
grazing pressure
Logging operations
and settlement
Excessive exploitation
Clearance for
agriculture, livestock
and plantation
development, timber
exploitation
Habitat destruction
and over-exploitation
Use as fuelwood,
fodder and browse
and changing land-use
patterns
Use as fuelwood and
grazing pressures
Over-grazing and
human pressure
Over-grazing and
human pressure
Utilisation of wood
and bark
Intensive agriculture
Intensive logging
Forest destruction and
exploitation for wood
and medicinal
purposes
Long-term climatic
changes; forest
clearance for
settlement and
cultivation, mineral
exploitation
Heavy exploitation,
plantation
development
Timber exploitation,
forest clearance for
pasture and
plantations
Table 25.3
FAMILY
Leguminosae (continued)
Meliaceae
Moraceae
Myrtaceae
Pinaceae
SPECIES
Mimosa
caesalpiniaefolia
Mimosa verrucosa
Pericopsis elata
Piptadenia peregrina
Plathymenia foliosa
Prosopis cineraria
Pterogyne nitens
StuhIlmannia moavi
Cedrela fissilis
Cedrela odorata
Entandrophragma
angolense
Khaya senegalensis
Lovoa swynnertonii
Milicia excelsa
Eucalyptus deglupta
Eucalyptus globulus
ssp. globulus
Abies guatemalensis
Abies nebrodensis
Abies numidica
Cedrus libani
DISTRIBUTION
Brazil
Brazil
West Africa to Zaire
Southern Brazil
Brazil
Arabia to India
Argentina, Brazil and
Paraguay
Tanzania
Costa Rica to
Argentina
Mexico to Argentina
and Caribbean Is
Tropical Africa
Tropical Africa
Tropical Africa
Tropical and sub-
tropical Africa
Philippines, Irian
Jaya and PNG
Southern Australia
and Tasmania
Mexico to El
Salvador
Sicily
Algeria
Lebanon and Turkey
347
Endangered tree species and provenances (continued)
STATUS
Suffering a slow
decline
Suffering a slow
decline
Endangered in parts
of its range and
subject to genetic
impoverishment
throughout
Threatened
Suffering a slow
decline
Endangered in parts
of its range
Becoming rare
Endangered
The best phenotypes
of many provenances
have mostly
disappeared
Endangered in parts
of its range
Threatened in parts of
W. Africa, severe
genetic erosion noted
in Nigeria
Populations of best
trees are in danger of
genetic erosion
Rare
Widespread but
threatened in some
areas
Endangered in parts
of its range
Threatened
Extremely rare and
threatened with
extinction
Endangered; about 20
wild trees remain
In danger of slow
decline
Threatened in
Lebanon
Plant Use
THREATS
Wood exploitation
Excessive exploitation
for the world timber
market and poor
natural regeneration
Clearance for
agriculture and cattle
rearing
Selective exploitation
Increasing human
pressure and changing
land-use patterns
Timber exploitation
Settlement and forest
clearance
Over-exploitation
Over-exploitation and
clearance of lowland
forest
Commercial
exploitation
Exploitation for timber
Forest clearance,
excessive
exploitation, poor
natural regeneration
Extensive logging
Limited regeneration,
logging and clearance
for agriculture
Development of pine
plantations
Illegal felling of small
Christmas trees
Exploitation and
increasing human
pressure
Grazing, local use of
wood
Exploitation and
grazing
2. Uses and Values of Biodiversity
Table 25.3
FAMILY
Pinaceae (continued)
Platanaceae
Rutaceae
Salicaceae
Simaroubaceae
Sterculiaceae
Taxodiaceae
Ulmaceae
Verbenaceae
SPECIES
Pinus armandii var.
amamiana
Pinus eldarica
Pinus koraiensis
Pinus patula ssp
tecunumanii
Pinus pentaphylla
Pinus pseudostrobus
Pinus radiata
Pseudotsuga
gaussenii
Pseudotsuga
sinensis
Platanus orientalis
Balfourodendron
riedelianum
Esenbeckia leiocarpa
Vepris glandulosa
Populus ilicifolia
Gymnostemon
zaizou
Nesogordonia
papaverifera
Glyptostrobus
lineatus
Taiwania
cryptomerioides
Taiwania flousiana
Ulmus wallichiana
Tectona
hamiltoniana
Tectona
philippinensis
DISTRIBUTION
Japan
USSR, Afghanistan
and Pakistan
Japan and the
Korean Peninsula
Central America
Japan and island of
Ullung-do (Korea)
Central America
California and
Mexico
Eastern China
China
E Mediterranean to
the Himalayas
Brazil, Paraguay and
Argentina
Brazil and Zaire
Kenya
Kenya
Cote d'Ivoire
West Africa
Widely cultivated in
parts of China, not
known in the wild
Taiwan and possibly
Myanmar
Myanmar, Tibet and
Yunnan
Afghanistan to
Nepal
Myanmar
Philippines
Endangered tree species and provenances (continued)
STATUS
Endangered
Endangered in the
USSR
In danger of depletion
in parts of its range
Under threat
throughout its entire
range
In danger of depletion
in parts of its range
Some provenances
are endangered
5 populations are
known; 1 is
endangered and the
genetic integrity of 2
others is endangered
Endangered
Naturally rare
Endangered in parts
of its range
Becoming scarce
Threatened in Brazil
Endangered
Endangered
Restricted distribution
Endangered in parts
of its range and
subject to genetic
impoverishment in
outlying populations
Natural populations
are extinct
Endangerd in certain
areas
Endangered
Endangered
Likely to be
endangered
Likely to be
endangered
THREATS
Lumbering
A relic species
Logging
Clearance for
agriculture and
attacks by Bark beetle
Logging operations
Selective logging
Grazing, urbanisation
and contamination by
cultivated stock
Agricultural expansion
and modification of
the water table
through irrigation
Habitat destruction
and exploitation
Forest clearance and
commercial felling
Settlement and forest
clearance
Habitat clearance
Logging
Large scale clear-
cuttings
Use as fodder
Local use for fuel and
construction and
forest fires
Naturally rare and
sought after for
general construction
Source: FAO 1986. Databook on Endangered Tree and Shrub Species and Provenances. FAO Forestry Paper 77. FAO, Rome.
348
Table 25.4
FAMILY
Combretaceae
Leguminosae
Meliaceae
Moraceae
Rubiaceae
Sapotaceae
Sterculiaceae
Plant Use
Commercial timber species of conservation concern in Ghana
SPECIES
Terminalia ivorensis
Terminalia superba
Guibourtia ehie
Pericopsis elata
Entandrophragma
angolense
Entandrophragma
candollei
Entandrophragma
cylindricum
Entandrophragma
utile
Guarea cedrata
Guarea thompsonii
Khaya anthotheca]/
grandifoliola
Khaya ivorensis
Lovoa trichilioides
Turraeanthus
africanus
Milicia excelsa
Milicia regia
Hallea ledermannii/
H. stipulosa
Nauclea didderichii
Aningeria robusta
Tieghemella heckelii
Mansonia altissima
Heritiera utilis
Nesogordonia
papaverifera
Triplochiton
scleroxylon
Source: Compiled from multiple sources.
TRADE NAME
Emire
Ofram
Anokye-hyedua
Kokrodua
(Afrormosia)
Edinam
Penkwa-akoa
(Candollei,Omu)
Penkwa (Sapele)
Efoobrodedwo
(Utile)
Kwabohoro
(Guarea)
Kwadwuma (Black
Guarea)
Krumben/Kruba
(Ahafo)
Dubini (Mahogany)
Dubinibiri (Walnut)
Apapaye (Avodire)
Odum
Subaha (Abura)
Kusia (Opepe)
Samfena (Aniegre,
Asanfona)
Baku (Makore)
Oprono (Mansonia)
Nyankom (Niangon)
Danta
Wawa
349
NO. OF
TREES
PER km?
45
13
12
18
14
5
152
EXPORT
OF
LUMBER
IN 1989
4697
32
10463
854
55
37747
combined
with
above
os
674
5668
778
1624
428
64818
CONSERVATION AND LEGAL
STATUS
Priority for in situ and ex situ
conservation. Threatened by over-
exploitation (FAO, 1984).
Priority for in situ and ex situ
conservation. Threatened by over-
exploitation (FAO, 1984).
ERL 18 years. Log export ban.
Threatened by over-exploitation
(FAO, 1984). Vulnerable. ERL O
years. Log export ban.
Priority for in situ conservation.
ERL 18 years. Log export ban.
Log export ban.
Priority for in situ conservation.
ERL 25 years. Log export ban
Priority for in situ conservation.
Threatened by over-exploitation
(FAO,1984). ERL 20 years. Log
export ban.
Priority for in situ conservation.
ERL Guarea spp. 82 years.
Priority for in situ conservation.
ERL Guarea spp. 82 years.
Priority for in situ and ex situ
conservation. ERL Khaya spp. 20
years. Log export ban.
Suffering from over-logging in
Ghana. Priority for in situ and ex
situ conservation.
Priority for in situ and ex situ
conservation. Threatened by over-
exploitation (FAO,1984)
Log export ban.
Priority for in situ conservation
Log export ban.
Priority for in situ conservation.
Vulnerable. ERL 10 years. Log
export ban.
Priority for in situ conservation.
Vulnerable. Log export ban.
H. ledermannii threatened by
over-exploitation (FAO, 1984).
Threatened by over-exploitation
(FAO, 1984).
Log export ban.
Log export ban.
Priority for in situ conservation.
Log export ban.
Log export ban.
Log export ban.
Priority for in situ and ex situ
conservation. Threatened by over-
exploitation (FAO,1984).
2. Uses and Values of Biodiversity
shows the species that are subject to a Ghanaian log export
ban. In addition to the log export ban, the Ghanaian
Government introduced Forest Improvement Levies in
November 1990. The highest rate of 50% is charged on
Nauclea diderrichii, 40% on Guarea cedrata and three
other species, and 10% on Triplochiton scleroxylon and one
other species. Levies on green/air-dried sawnwood were:
50% on Pericopsis elata, Entandrophragma utile, Guibortia
ehie and Tieghemella heckelii and 8.5% on Milicia excelsa
and M. regia. The Government plans to follow up the levies
by introducing a ban on exports of green sawn timber in
January 1994.
In addition to the species that have suffered genetic erosion
because of heavy exploitation, there are many rare timber
species in Ghana that are not currently recorded in the
timber export trade. A recent field guide to the forest trees
(Hawthorne, 1990), for example, notes 27 timber species as
being uncommon, rare, or very rare. Some of these are of
more immediate conservation concern than the major
commercial species. Overall the most serious threat to tree
species in Ghana comes from fire damage, which has
severely undermined the regeneration of trees even in the
most productive moist semi-deciduous forest zone.
RATTANS
After timber, rattans (lianoid palms) provide the second
most important source of export earnings from tropical
forests. Most of the 600 or so species are native to South
and Southeast Asia. Countries with major rattan industries
include the Philippines, China, Indonesia, India, Sri Lanka
and Thailand, and these provide full-time employment for
at least half a million people. For the international market,
rattans are mainly used in the production of cane furniture.
Local uses include the production of mats, baskets, fish
traps, dyes and medicines.
The rattan industry relies almost entirely on wild stocks.
About 90% of the world’s raw material supply is extracted
from the wild and the remaining 10% from plantations in
Central and South Kalimantan. Exploitation combined with
habitat destruction has led to the decline of major
commercial rattan species and species that are valuable in
local use and local markets. Table 25.5 lists the major
commercial rattan species with notes on their conservation
Status.
Indonesia is the world’s main producer of rattans, supplying
about 90% of the total raw material utilised. Export of raw
rattan from the country has been banned since 1979. There
has been relatively little downstream processing of rattans
into finished products within Indonesia and, in an attempt
to boost local value-added production, a ban on export of
non-finished products was introduced in 1989. This has led
to concern about increasing commercial pressure on wild
stocks elsewhere.
The centre of diversity for rattans is the Malay Peninsula.
A total of 104 species occur within this area, of which
about 38% are endemic. Of these Malay Peninsula species,
only two are considered to be not threatened and 98 are
categorised as Vulnerable or Endangered (Kiew and
Dransfield, 1987). Research has begun on the taxonomy and
350
silviculture of Malaysian rattans as a prelude to bringing
these into cultivation. In the meantime, it is uncertain how
many of the 104 species in Peninsular Malaysia occur
within the State’s existing national park (Taman Negara).
Illegal removal of commercial species remains a threat
within the protected area. Ex situ conservation of rattan
species in seed banks is not currently a viable proposition
because rattans have recalcitrant seeds. The most attractive
form of genetic conservation for rattans in Peninsular
Malaysia will probably be through their cultivation in
logged-over hill dipterocarp forests.
MEDICINAL PLANTS
Around 119 pure chemical substances extracted from some
90 species of higher plants are used in medicines throughout
the world. At a local level an extremely wide range of plant
species is used medicinally. The World Health Organization
has listed over 21,000 plant names (including synonyms)
that have reported medical uses around the world. Very few
of these medicinal plants have been subject to scientific
scrutiny. In all about 5,000 higher plant species have been
thoroughly investigated as potential sources of new drugs.
Most of these are temperate species and the biochemical
potential of tropical plants has been largely overlooked.
Nevertheless around 80% of people in developing countries
rely on traditional medicines. Table 25.6 shows some of the
most important plant species whose derivatives are used in
orthodox medicine along with an indication of whether
analogous uses have been reported in traditional medicine.
Medicinal plant species are still to a large extent harvested
from the wild and relatively few are cultivated as crop
plants. For example in Germany two-thirds of the species
used are still wild collected and cultivation of major
medicinal plants such as Gentiana lutea, Valeriana
mexicana, Echinacea and Arnica has only begun in the past
20 years. Plant breeding has only taken place with the
commercially most important plants such as Papaver
somniferum, Papaver bracteatum, Cinchone pp.,
Chamomilla recutita and Mentha piperita (Schumacher,
1991). In many cases, biochemicals extracted from plants
have been used as blueprints for the synthesis of drugs and
the natural source material is no longer required.
Nevertheless, the USA annually imports over US$20
million worth of rain forest plants for medicinal purposes.
Important drugs include tubocuranin, derived from plant-
based curare and used as a muscle relaxant during surgery,
and curianol, a Guyanese fish poison used in heart
operations. Economic aspects of the production of
pharmaceuticals from plants are discussed in Chapter 27.
The US National Cancer Institute has identified over 1,400
tropical forest plants with the potential to fight cancer. One
such plant is the Rosy Periwinkle Catharanthus roseus
native to Madagascar. Used for generations by tribal
healers, this species is now used in the production of drugs
effective against Hodgkins disease and other forms of
cancer. The Rosy Periwinkle yields vinca alkaloids, which
are complex molecules difficult to synthesise chemically. It
remains cheaper to collect leaves of living plants for
extraction of the valuable medicinal products. Catharanthus
roseus is now a widespread weedy species in the tropics and
is commonly cultivated. All other species of the genus are
Table 25.5
SPECIES
Calamus caesius Blume
Calamus diepenhorstii Miq.
Calamus manan Miq
Calamus maximus Merr.
Calamus mindorensis Becc.
Calamus optimus Becc.
Calamus ornatus Bl.
Calamus peregrinus Furtado
Calamus rudentum Lour.
Calamus scipionum Lour.
Calamus subinermis H.
Wendl.
Calamus trachycoleus Becc.
Calamus tumidus Furtado
RANGE
Malay Peninsula, Borneo, Sumatra,
Philippines (Palawan); Thailand (possibly
introduced)
Malay Peninsula, Singapore, Sumatra,
Borneo (Sabah), Philippines (Palawan)
Malay Peninsula, Borneo, Sumatra, south
Thailand
Philippines (Basilan, Luzon, Mindanao,
Mindoro)
Philippines (Luzon, Mindoro)
Borneo
Malay Peninsula, Borneo, Sumatra,
Sulawesi, south Thailand, Philippines
(Luzon, Mindanao, Mindoro, Negros,
Palawan, Polilo)
Malay Peninsula, Thailand
Thailand, Laos, Cambodia, Viet Nam
Malay Peninsula, Singapore, Borneo,
Sumatra, Philippines (Palawan)
Sabah
Kalimantan
Malay Peninsula, Sumatra
Plant Use
Main commercial species of Rattan (Palmae: Ca/amus)
STATUS AND THREATS
Domesticated in Kalimantan. Supply of wild
stocks threatened by over-exploitation
Threatened: viable populations largely limited
to a few inaccessible areas as a result of
excessive and premature exploitation
Any accessible populations have been
exploited; but the species clusters and so is
not as vulnerable to over-exploitation as is
(say) C. manan. However, there has been
extensive habitat destruction by logging,
shifting cultivation and spontaneous
settlement
*
Endangered: a rare and much sought after
species; so much so that it is very difficult to
find mature long canes even in Mulu National
Park (Sarawak)
*
*
Extensive habitat destruction
*
Present stocks are limited and their
exploitation requires strict control
Domesticated in Kalimantan
Largest known populations threatened by
Calamus zollingeri Becc. Sulawesi
agriculture
*
Source: Dransfield, J. 1979a. A Manual of the Rattans of the Malay Peninsula. Malayan Forest Records 29. Forest Department. Ministry of
Primary Industries, Malaysia. Dransfield, J. 1979b. Report of Consultancy on Rattan Development carried out in Thailand, Philippines, Indonesia
and Malaysia. 14 March-8 May 1979. For FAO Regional Office for Asia and the Far East, Bangkok. Dransfield, J. 1981. The biology of Asiatic
Tattans in relation to the rattan trade and conservation. In: Synge, H. (Ed.), The Biological Aspects of Rare Plant Conservation. Wiley, Chichester.
Dransfield pers. comm. 18 February 1981.
Notes: * No information.
endemic to Madagascar, where several are used
medicinally. One species which has not been tested
phytochemically is close to extinction.
Regions that are known to have important concentrations of
major medicinal plants include Mexico and Central
America, the west-central region of South America
(Colombia, Ecuador and Peru), the Indian subcontinent,
west Asia and parts of north-eastern Africa. Over-
exploitation of medicinal plants extracted from the wild is
leading to problems of genetic erosion in some of these
regions. In India, for example, where 2,500 plant species
are used by traditional healers, species of Aconitum,
Dioscorea and Ephedra are some of the medicinal plants
under threat in the wild.
Dioscorea deltoidea, a species that grows in the Himalayan
foothills of northern India, is a major source of diosgenin
351
used in the manufacture of contraceptive pills. Over-
collection has led to the decline of this species in the wild
and it is now subject to international trade controls. The
remaining small specimens of the plant yield less than 15%
of the diosgenin found in the large, old tubers which have
mostly been removed from the wild.
Also threatened in India is Rauvolfia serpentina, a forest
shrub known as serpentine root. This has been used in
traditional medicine for 4,000 years to treat snakebite,
nervous disorders, dysentery, cholera and fever. An extract
from the plant, reserpine, became the principal source of
materials for modern tranquilisers following research
around 50 years ago. In Thailand, Rauvolfia serpentina is
collected both for use in local medicine and for sale, via
middlemen, to national and international pharmaceutical
companies.
2. Uses and Values of Biodiversity
Table 25.6
Principal plant species with constituent compounds used as drugs
PLANT NAME COMPOUND NAME THERAPEUTIC CATEGORY PLANT USES CORREL- COUNTRY OF PRODUCTION OR
IN MEDICAL SCIENCE IN ATION CULTIVATION
TRADITIONAL BETWEEN
MEDICINE TWO USES
Ammi spp. Xanthotoxin Pigmenting agent Leukoderma; Yes Cultivated in Asia and the
Vitiligo Mediterranean region
Khellin Bronchodilator Asthma Yes
Atropa Atropine Anticholinergic Dilate pupilof Yes Central and Southern Europe,
belladonna eye cultivated in USA, UK, Easter India,
Europe, China
Berberis vulgaris Berberine Antibacterial Gastric Yes Europe, Asia
ailments
Carica papaya Chymopapain Proteolytic; mucolytic Digestant Yes Cultivated in Sri Lanka, Zaire,
Papain Proteolytic; mucolytic Digestant Yes Uganda, Mozambique, Tanzania,
South Africa, India
Cassia spp. Danthron Laxative Laxative Yes C. acutifolia cultivated in India
C. senna cultivated in Egypt
Catharanthus Vinblastine Antitumor agent Not used No Pantropical, cultivated in US, India
roseus Vincristine Antitumor agent Not used No and other countries
Cephaelis Emetine Amebicide; emetic Amebicide; Yes Brazil, much collected in Mato
jpecacuanha emetic Grosso
Cinchona Quinidine Antiarrhythmic Malaria No Cultivated in Indonesia, Zaire,
ledgeriana Quinine Antimalarial; antipyretic Malaria Yes Tanzania, Burundi, India, Kenya,
Guatemala, Peru, Ecuador, Bolivia,
Rwanda, Sri Lanka, Colombia, Costa
Rica
Datura matel Scopolamine Sedative Sedative Yes Cultivated in Asia
Digitalis spp. Acetyldigitoxin Cardiotonic Not used Indirect D. lanata Cultivated in southem
Deslanoside Cardiotonic Not used Indirect Europe and Asia, D. purpurea
Digitoxin Cardiotonic Cardiotonic Yes cultivated in India and temperate
Lanatosides Cardiotonic Not used Indirect zones
Digitalin Cardiotonic Cardiotonic Yes
Gitalin Cardiotonic Cardiotonic Yes
Ephedra sinica Ephedrine Sympathomimetic Chronic Yes China
bronchitis
Pseudoephedrine* Bronchodilator Chronic Yes
bronchitis
Glycyrrhiza Glycyrrhizin Sweetener Sweetener Yes Cultivated in Spain, Turkey, Iraq,
glabra (Glycyrrhetic acid) China, Mongolia, USSR, South
Africa, USA, France, Italy, Iran,
Afghanistan, Syria, Lebanon, Israel,
UK, China
Hyoscyamus Hyoscyamine Anticholinergic Sedative Yes Cultivated in temperate zones
niger
Papaver Codeine Analgesic; antitussive Analgesic; Yes Cultivated in Turkey, India, Burma,
somniferum sedative Thailand
Morphine Analgesic Analgesic; Yes
sedative
Noscapine (narcotine) Antitussive Analgesic; Yes
Papaverine* sedative
Smooth muscle relaxant Sedative; No
analgesic
Pausinystalia Yohimbine Adrenergic blocker; Aphrodisiac Yes Cameroon, Nigeria, Rwanda
yohimbe aphrodisiac
Physostigma Physostigmine Anticholinesterase Ordeal poison Indirect Sierra Leone, Cameroon, introduced
venenosum (eserine) to India and Brazil
Pilocarpus Pilocarpine Parasympathomimetic Poison Indirect Tropical America especially Brazil
jaborandi
Rauvolfia spp. Ajmalicine Circulatory stimulant Tranquilizer Indirect e.g. A. serpintina; Thailand, Zaire,
Rescinnamine Antihypertensive; Tranquilizer Yes India, Bangledash, Sri Lanka, Burma,
tranquilizer Malaysia, Indonesia, Nepal
Reserpine Antihypertensive; Tranquilizer Yes
tranquilizer
Silybum Silymarin Antihepatotoxic Liver disorders Yes Mediterranean region
marianum
Urginea Scillaren A Cardiotonic Cardiotonic Yes Mediterranean region, Egypt, Turkey
maritima
Valeriana Valepotriates Sedative Sedative Yes Cultivated in Eastern Europe,
officinalis Netherlands, Japan
Note: * Also now synthesised commercially. Source: Farnsworth, N.R. 1988. Screening plants for new medicines. In: Wilson, E.O. (Ed.),
Biodiversity. National Academy Press, Washington. Pp.83-97; Farnsworth, N.R. and Soejarto, D.D. 1991. Global importance of medicinal plants.
In: Akerele, O., Heywood, V. and Synge, H. (Eds), The Conservation of Medicinal Plants. Proceedings of an International Consultation 21-27
March 1988. Chiang Mai, Thailand. Cambridge University Press, Cambridge, UK.
352
In many parts of the tropical world there is a serious lack
of knowledge about the genetic resources and conservation
status of the medicinal plants on which most people rely. In
Malaysia, for example, the genetic resources of medicinal
plants have scarcely been evaluated. At present although
some medicinal plants are cultivated on a small scale, most
herb traders and local medicine men rely on wild resources.
There is some concern that collectors do not know the
status of individual species and may contribute to the loss
of populations of threatened species. Clearance of
Malaysian forests is likely to lead to the loss of medicinal
plant species in families such as Apocynaceae, Annonaceae,
Rutaceae, Dioscoreaceae, Leguminosae, Lauraceae and
Zingiberaceae. Unlike the fruit tree resources, which have
survived under semi-wild cultivation in village orchards,
medicinal plant resources will be lost from forest areas.
In Thailand most of the plants used in rural medicines are
collected from forests. Many of these have commercial
value as raw materials used by over 1,000 traditional drug
manufacturing companies, modern drug companies and in
export as crude extracts. Thailand is expanding its domestic
production of herbal medicines but increasingly depends on
imported raw materials as local resources are lost.
The use of plant resources in Indonesia for the production
of ‘jamu’ (herbal medicine) has generally been sustainable
for home consumption. Increasing commercialisation is,
however, putting pressure on wild populations of medicinal
plants. Modern jamu industries and pharmaceutical
companies are using large quantities of plant materials and
this is leading to genetic erosion of species that are not in
cultivation. Species that are being affected include Curcuma
spp., which were previously abundant in East Java but now
have to be imported from other islands.
Another species, Parkia roxburghii, has also become rare
in parts of East Java, owing to excessive harvesting of the
fruits. Increasing international demand for Curcuma spp.,
and others such as Voacanga gradifolia, Orthosiphon
aristasus and Rauvolfia, is leading to the loss of
economically valuable plant genetic resources that could
provide for a sustainable source of foreign exchange if
brought into cultivation.
One of the few medicinal plants which has been developed
as a major crop species is quinine Cinchona spp. The main
use of quinine, extracted from bark of the Cinchona trees,
is in anti-malarial drugs. Initially the whole world supply
came from wild trees in the Andes. This led to concern
about the possible extinction of the species and as a result,
plantations were developed in the middle of the last century,
for example in India and Indonesia. Synthetic alternatives
are available but quinine remains an important drug with
new applications being found. The genetic base of the crop
is very narrow and conservation of wild stands of Cinchona
is important for future breeding work.
ORNAMENTAL PLANTS
The discovery, domestication and cultivation of ornamental
plants have a long history, comparable to that of food
crops. Lilies, for example, have been cultivated in China
353
Plant Use
for both medicinal and decorative purposes for around two
thousand years. In Roman times, roses, lilies, violets,
anemones, narcissi and lavender were grown as garden
plants in Europe. Today, the diversity of decorative plant
species established in cultivation far surpasses the variety of
plants commonly grown for food around the world. In the
UK alone, an estimated 3,000 species are in general
cultivation in addition to the wide range of cultivars and
hybrids. At least five times as many species have been
introduced at various times in the past. Novelty and variety
remain important factors in the horticultural market.
Ornamental plants are an important commodity in
international trade, with an expanding international market.
Total world imports of cut flowers, cut foliage and plants
amounted to US$2,488 million in 1985. The value of world
trade in cut flowers and live plants for 1981 to 1985 is
shown in Table 25.7. It should be stressed that virtually all
this value is in artificially-propagated stock.
Despite the economic importance of ornamental plants, the
conservation of these genetic resources is usually given a
low priority both nationally and internationally when
compared to food, fruit and forage crops (Chin, 1989).
Wild species of horticultural value are under threat around
the world, both through the processes of habitat destruction
and through direct exploitation for local use and
international trade.
Although sophisticated propagation techniques have been
developed for ornamental plants, significant quantities of
plants in some groups continue to be dug from the wild for
the world market. This is apparent for example with bulbs,
orchids, cacti and other succulent plants, cycads and
insectivorous plant species. Concern about the level of
exploitation of some of these plants has led to their listing
on the Appendices of CITES. As a result, data on levels of
international trade in both wild-collected and artificially
propagated plants have been recorded. Summary figures for
cactus and orchid trade for 1989 are given in Table 25.8.
Orchids
Over 5,000 orchid species have been recorded in CITES
trade statistics during the period 1983-1989, with the
average annual number of plants in international trade being
nearly five million. This figure excludes orchids in flasks
and cut flowers. Around 80% of the orchids in trade are
reported to be artificially propagated and most trade is in
artificially propagated hybrids. There is still, however,
significant international demand for species orchids and a
large part of this trade is satisfied through the collection and
export of wild plants. The major source country for orchids
in international trade is Thailand. Orchids are propagated in
commercial nurseries within the country but, at the same
time, there is a huge trade in both native and imported wild
orchids.
The most heavily traded orchid genus exported from
Thailand is Dendrobium. The numbers of native species
exported together with the levels of plants reported to be
propagated are shown in Table 25.9.
2. Uses and Values of Biodiversity
Table 25.7 Value of world trade in flowers and plants, 1981 —1985,
(US$ millions)
CUT FLOWERS
IMPORTS ANNUAL MEAN %
1238.79
EEC
Germany, Fed Rep 535.54 43.23
France 85.55 6.91
United Kingdom 70.27 5.67
Netherlands 53.82 4.34
Italy 25.49 2.06
Belgium—Luxemburg 29.33 2.37
Denmark 11.28 0.91
Ireland 3.48 0.28
Greece 0.41 0.03
United States 206.90 16.70
Canada 24.17 1.95
Japan 18.08 1.46
Switzerland 63.14 5.10
Austria 37.81 3.05
Sweden 30.19 2.44
Norway 12.05 0.97
Singapore 5.07 0.41
Finland 6.38 0.52
Hong Kong 5.21 0.42
Saudi Arabia 3.58 0.29
Australia 2.65 0.21
Kuwait 1.16 0.09
United Arab Emirates 1.01 0.08
Spain 0.56 0.05
LIVE PLANTS
IMPORTS 915.76
EEC
Germany, Fed Rep 228.64 24.97
France 116.20 12.69
United Kingdom 80.00 8.74
Italy 57.14 6.24
Netherlands 50.99 5.57
Belgium—Luxemburg 41.72 4.56
Denmark 13.94 1.52
Greece 4.36 0.48
Ireland 3.99 0.44
United States 37.67 4.11
Canada 43.69 477
Japan 7.15 0.78
Sweden 60.03 6.56
Switzerland 41.91 4.58
Austria 20.99 2.29
USSR 21.91 2.39
Spain 16.52 1.80
Finland 14.07 1.54
Norway 12.24 1:34
Saudi Arabia 7.65 0.84
Colombia 4.20 0.46
Hong Kong 2.77 0.30
Algeria 2.23 0.24
Singapore 1.82 0.20
EXPORTS ANNUAL MEAN %
1101.79
Netherlands 701.51 63.67
Colombia 121.68 11.04
Israel 73.31 6.65
Italy 73.35 6.66
Spain 17.96 1.63
Thailand 16.65 1.51
France 14.03 1.27
Kenya 7.60 0.69
Taiwan Province (China) 6.28 0.57
Germany, Fed Rep 5.59 0.51
United States 9.92 0.90
South Africa 6.17 0.56
Singapore 6.83 0.62
United Kingdom 4.36 0.40
Peru 3.18 0.29
New Zealand 3.03 0.28
Mexico 3.49 0.32
Costa Rica 1.29 0.12
Brazil 2.31 0.21
Ethiopia 0.86 0.08
Morocco 1.41 0.10
Malaysia 1.34 0.12
Mauritius 0.70 0.06
882.15
Netherlands 389.70 44.18
Denmark 123.77 14.03
Belgium—Luxemburg 92.62 10.50
Germany, Fed Rep 51.86 5.88
France 37.30 4.23
United States 33.64 3.81
Italy 26.94 3.05
Canada 18.16 2.06
Spain 16.08 1.82
Guatemala 10.00 1.13
Costa Rica 8.31 0.94
Japan 6.20 0.70
Israel 5.32 0.60
United Kingdom 4.39 0.50
Cote d'Ivoire 4.15 0.47
New Zealand 2.49 0.28
Honduras 3.36 0.38
Hungary 1.89 0.21
Brazil 1.77 0.20
Singapore 2.08 0.24
Malaysia 0.73 0.08
Egypt 1.41 0.16
Jamaica 1.05 0.12
Thailand 0.78 0.09
Colombia 1.56 0.18
Turkey 0.79 0.09
Source: International Trade Centre UNCTAD/GATT. 1987. Floricultural products: a study of major markets. Genera.
The impact of orchid collection within Thailand has been
highly detrimental. The conservation status of native species
is scarcely known, but it is apparent that some species have
been virtually eradicated even within national parks where
collection is banned. Increasing prices reflect the increased
scarcity of desirable species.
Orchid collection, together with habitat destruction, has led
to the decline of wild orchid species in many other
countries. In Japan, for example, more than 70 orchid taxa
354
are included in the Japanese Plant Red Data List, of which
50 are threatened by over-collection. The genus Calanthe
has been particularly popular with collectors and is under
great pressure in the wild. Other genera which have been
seriously over-collected include the Asian slipper orchids in
the genus Paphiopedilum and the Latin American slipper
orchids in the genus Phragmipedium. Both these genera are
now included in Appendix I of CITES which effectively
bans commercial international trade in wild-collected
specimens.
Plant Use
Table 25.8 Cactus and orchid trade data for 1989
cACcTI ORCHIDS
IMPORTS' EXPoRTsS' IMPORTS! EXPORTS!
WORLD 6513647 6513647 8313088 8313088
ASIA: 209716 3749060 6017522 7133797.
Brunei 0 0 2388 0
China 200 te) ) 165505
Cyprus 17801 t) 356 0
Hong Kong 45313 0 114841 ()
India 3 i) ie) 8423
Indonesia 6 (0) 4888 0
Israel te) (0) 1660 0
Japan te) 1708096 5509995 (1)
Korea, Dem People’s Rep (0) (0) 1100 0
Korea, Rep _ (0) 2040964 381027 (t)
Macau 0 0 21 0
Malaysia 9000 0 te) 22117
Nepal 1105 {0} 0 16
Pakistan 3 ie) te) 0
Philippines (0) (0) 0 50971
Saudi Arabia 1371 0 40 0
Singapore 56204 0 0 27275
Sri Lanka (0) 0 0 20283
Syria 0 0 5 ts)
Taiwan 71464 0 {0} 1594732
Thailand 4 0 0 5244450
Turkey 330 0 52 0
United Arab Emirates 6912 0 1149 (:)
Viet Nam 0 0 (0) 25
USSR* 0 0 0 1499:
EUROPE 3038304 100906 1671781 984041
Austria 195652 te) 67590 (0)
Belgium 0 6494 8077 0
Czechoslovakia te) 433 2427 0
Denmark 6533 (0) 1753 (0)
Finland 44136 0 4942 0
France 0 70991 9600 0
German Dem Rep 2 0 2519 0
Germany, Fed Rep 189264 0 1138720 0
Hungary 0 (0) 4732 0
Iceland 0 0 14 tt)
Ireland (0) 0 25 0
Italy 28138 0 14841 0
Luxembourg 0 (0) 456 0
Malta 258 to) 388 (0)
Monaco (0) 3 (0) ()
Netherlands 1991664 0 305494 0
Norway 12956 (0) 131 (1)
Poland te) (¢) (0) 24450
Portugal to) 0 110 0
Spain (0) 22985 154 it}
Sweden 165480 14431 0
Switzerland 382916 0 95377 0
United Kingdom 18153 0 0 959591
Yugoslavia 3152 0 0 0
NORTH & CENTRAL AMERICA 3204234 1462004 408654 40126
Antigua 0 0 81 0
Aruba (e) 8 203 0
Bahamas 1054 0 1683 ts)
Barbados 1594 (0) 259 t)
Belize 0 6 (0) 4387
Bermuda 0 0 4098 0
Canada te) 1032492 118474 te)
Cayman Islands 0 (0) 978 0
Costa Rica (0) (0) 3499 i)
Dominica te) (0) (0) 180
Dominican Rep 0 347857 692 0
El Salvador 0 (0) 197 (Y)
Guadeloupe (0) 0 45 0
Guatemala 0 4 0 69
Haiti (0) 3739 734 0
Honduras 0 1 (0) 13205
Jamaica (e) 1 () 1419
Martinique 0 0 576 tt)
Mexico 0 77896 0 20716
Montserrat (0) (0) 760 tt)
Netherlands Antilles 0 0 1 0
Nicaragua 0 0 0 150
Panama 300 (0) 15982 te)
Puerto Rico (0) 0 221 te)
St Lucia 300 0 318 te)
Trinidad and Tobago 1495 0 11494 0
United States 3199272 0 248359 0
Virgin Islands (British) 219 (0) (0) 0
SOUTH AMERICA 17 1127863 5203 124525
Argentina 0 396 9 ts)
Bolivia te) 0 20 t)
Brazil (e) 1127181 0 93426
Chile 0 59 68 ts)
Colombia 0 17 ie} 8965
355
2. Uses and Values of Biodiversity
Table 25.8 Cactus and orchid trade data for 1989 (continued)
CACTI ORCHIDS
IMPORTS! EXPoRTs' IMPORTS! Exports!
SOUTH AMERICA (continued)
Ecuador ts) 0 (0) 795
Guyana 0 0 5030 t!)
Paraguay (0) 7 0 872
Peru te) 186 te) 18078
Suriname () 0 76 i}
Uruguay 17 () [*) 0
Venezuela 0 17 0 2389
OCEANIA 60327 0 138112 11023
Australia 56632 0 71134 0
Fiji te) 0 23837 (9)
French Polynesia 0 t) 41286 0
New Caledonia ts) 0 1811 0
New Zealand 3695 0 0 8149
Palau 0 0 44 0
Papua New Guinea 0 0 i) 2668
Vanuatu 0 0 te) 206
AFRICA 359 73807 56273 18040
Botswana 0 0 21 0
Cameroon 76 0 46 t)
Comoros te) 0 te) 50
Céte d'ivoire te) 0 te) 526
Ethiopia (e) () (e) 66
Gabon 0 (0) 1017 ()
Ghana te) (°) (0) 2
Kenya (0) 0 i) 4819
Liberia fe) (e) 0 6
Madagascar 0 3 i*} 12459
Mauritius te) 0) 1271 ts)
Morocco 0 51607 41 1)
Mozambique i) 0 50 0
Nigeria (*) te) te) 9
Reunion 60 i} 4718 0
South Africa 0 22197 49072 0
Togo 2 0 0 t)
Tunisia te) (e) 8 ()
Zaire 0 0 14 0
Zambia () () 15 te)
Zimbabwe 221 0 (0) 103
OTHER 690 7 15543 37
Country Unknown 690 0 15527 0
Other (0) 7 16 37
Notes: | Figures are net. * Former USSR.
Sources: Annual reports of Parties to CITES compiled by WCMC.
Table 25.9 Dendrobium Orchids from
Thailand
SPECIES AVERAGE NO. % ARTIFICIALLY
IN ANNUAL TRADE PROPAGATED
1983-1989
D. aphyllum # 740 11
D. bellatulum # 1526 19
D. chrysotoxum # 5110 25
D. densiflorum * 777 30
D. draconis 985 13
D. farmeri # 2396 18
D. fimbriatum 464 16
D. harveyanum 1022 14
D. nobile 812 90
D. parishii 1960 45
D. scabrilingue # 1779 11
D. senile # 1160 30
D. thrysiflorum # 2959 27:
D. unicum # 1175 4
Notes: # Known to be sold as wild-collected plants in Thailand.
* Doubtfully native to Thailand.
Source: Oldfield, S. 1991. Review of significant trade in species of
plants included in Appendix II of CITES, 1983-1989. Report prepared
for the 8th meeting of the Conference of Parties. Unpublished Report.
356
Cacti and other succulents
The average annual international trade in cacti as recorded
in CITES statistics is close to 14 million. This is probably
an underestimate of the real trade, because a single nursery
in the Netherlands (which exports most of its production)
produces over 18 million cacti annually and commercial
cactus production in the USA has been estimated at 10-50
million per year. The bulk of cacti in international trade are
propagated but collection continues to put pressure on
certain desirable species which are close to extinction in the
wild. Mexico, one of the main centres of diversity of the
cactus family, exports around 50,000 cacti annually
according to CITES figures. A high proportion of these are
wild-collected and exported illegally.
A wide range of other succulent plants, including species of
Aloe, Euphorbia and Pachypodium, are also traded
internationally. One of the main source countries is
Madagascar which has exported around 135,000 CITES-
listed succulents annually, all of which are wild-collected.
This trade poses a severe threat to Madagascar’s unique
plants and adds to the pressures of habitat destruction.
Plant Use
Table 25.10 Trade and conservation status of Turkish bulbous species
CONSERVATION
CATEGORY
GENUS/SPECIES
Allium roseum
Anemone blanda
Arum spp.
Crocus spp.
Cyclamen spp.
Cyclamen cilicium
C. graceum
C. hederifolium
C. mirabile
C. persicum
Cs, ICS CC CC Ii,
C. repandum
Dracunculus spp.
<
Eranthis hyemalis
Fritillaria imperialis
m
F. persica
Galanthus spp.
Hyacinthus orientalis orientalis
Leucojum aestivum
Lilium candidum
mmm<«<cdc
L. martagon
Muscari spp.
Narcissus spp.
N. serotinus
Pancratium maritium
Scilla spp.
Sternbergia spp.
Tulipa spp.
T. praecox
T. humilis
Urginea maritima
CULTIVATION IN
EXPORT EXPORT TURKEY (WHERE
FIGURES (1987) CONTROLS KNOWN)
B
7,500,000 Q
Q
B
995,000 Q
wild-transplanted
Q
10,000,000 Q artificial propagation
275,000 B artificial propagation
275,000 B wild-transplanted
30,000,000
(G. elwesii and G.
ikariae) Q
B
8,500,000 artificial propagation
1,335 wild-transplanted
artificial propagation
B/Q
Q
B
100,000
450,000
(S. Jutea (V) and
S. clusiana) wild transplanted
B
37,000 Q
Sources: Ekim, T., Koyuncu, M., Erik, S. and Darslan, R. 1989. List of Rare, Threatened and Endemic Plants in Turkey, prepared according
to IUCN Red Data Book categories. Turkish Association for Conservation of Nature and Natural Resources, Ankara. Series No. 18. McGough,
H.N., Mathew, B.F., Peter, H., Read, M., Wertel N. and Wijnands, O. 1989. A report on the status and cultivation of Cyclamen species and other
geophytes in Turkey. Paper prepared for the Scientific Working Group of the EC CITES Committee.
Notes: E Endangered; V Vulnerable; R Rare; I Indeterminate; B Ban on export; Q Quota system for exports
Bulbs
Information on levels of international trade in wild bulbs is
less readily available because most genera are not covered
by CITES. Commercial cultivation of most bulbous genera
is well-established but collection from the wild takes place
routinely for certain so-called minor bulbs such as
snowdrops (Galanthus) and Cyclamen. In general, it is
difficult to assess the impact of collection on wild bulb
populations but genetic erosion is a serious problem for
species of horticulturally popular genera.
The daffodil genus Narcissus has around 40 species, with
its centre of diversity in Spain and Portugal. Hundreds of
daffodil cultivars and hybrids have been developed and
357
daffodils are an important horticultural crop in various
countries. The UK is the major exporter of daffodil bulbs,
with five varieties dominating commercial production. In
1987 the UK exported nearly 87 million Narcissus bulbs
with a value of over £4 million. At present there is very
limited UK production of small Narcissus species which are
becoming increasingly popular. One of the main sources of
these species is Portugal where the bulbs are dug from the
wild.
Ten taxa of Narcissus are considered to be threatened in
Portugal and to be in need of protection. Several of these,
including N. asturiensis and N. cyclamineus, are exported
to the Netherlands for re-export around the world.
2. Uses and Values of Biodiversity
The main source country for wild-collected bulbs in
international trade is Turkey. The country has a very rich
bulbous plant flora and is the origin of many of the
attractive bulbs in cultivation. Commercial exports of bulbs
from Turkey are subject to licensing by the Turkish
Government and official statistics are based on the
quantities licensed. Turkish exports of the main commercial
genera are given in Table 25.10 above.
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Chapter provided by Sara Oldfield.
26. ANIMAL USE
INTRODUCTION
Wildlife can be used in a variety of ways, involving
different degrees of human intervention and modification of
natural habitats, with varying effects on conservation. At
one end of the spectrum lie in situ harvesting regimes, such
as the harvesting of wild plants and animals for subsistence
use by local communities. Many, perhaps most, harvests of
wild species have in the past resulted in population declines
but, if practised at appropriate intensity, these forms of use
could be sustainable and need entail little alteration of
natural ecological processes. In such cases they might be of
conservation benefit if they also provided an economic
incentive for conserving natural habitats. At the other end
of the scale, wildlife can be brought into captivity and
reared in a controlled environment (a process leading
eventually to domestication) which usually has little or no
conservation value. In between these two extremes there is
a range of different containment or husbandry systems,
varying from the intensive to the extensive. The
implications for the conservation of biodiversity of these
different forms of use are considered below.
Many of the products obtained from wildlife are exploited
commercially. Worldwide, the commercial trade in wild
plants and animals was valued by Hemley (1988) at US$5
billion. Many of the arguments used to justify the
conservation of biodiversity rely on the benefits that can be
obtained, both economic and otherwise, from the
sustainable use of wild resources. However, it should be
stressed that, whatever the management system involved,
arguments for the conservation of wildlife on purely
economic grounds may be insufficient to ensure its long-
term preservation. If a landowner is convinced that wildlife
should be conserved because of its profitability compared to
other forms of land-use, then the logical extension of the
argument is that if it ceases to be profitable he should
remove it. It should therefore be remembered that aesthetic
and moral grounds are just as valid as financial arguments
as justifications for the conservation of wildlife.
FOOD: TERRESTRIAL ANIMALS
Vertebrates
Although most of the human diet is now more generally
provided by domesticated animals and plants, and fisheries,
other wildlife still feature as an important source of
nutrition (Table 26.1). Information on wildlife consumption
is sparse and typically non-quantitative, partly because of
the nature of the consumption, which is generally on a
subsistence basis and therefore unrecorded by the normal
accounting processes. Estimated figures vary wildly and are
often contradictory. However, case-studies reveal the
pervasive nature of subsistence-based wildlife use in many
cultures and societies. For example, in the Huallaga Central
Region of Peru, new settlers and the indigenous peoples
Tely on wildlife for as much as 80% of their animal protein
(Library of Congress, 1979). In northern Alaska in 1974,
the people of the Anaktuvuk Pass each consumed an
average of 755kg of meat from wild animals, some 88% of
359
Animal Use
their diet. By 1984, this proportion had fallen to 70%
(Klein, 1989). The Mbuti pygmies of Zaire obtain up to
60% of their calorific intake from hunting (Marks, 1989).
The majority of animal protein consumed by rural
communities around Kisangani (Zaire) derives from wild
animals, mostly duikers, rodents, primates and other small
mammals such as bush pigs, bats and pangolins (Colyn et
al., 1988). In Liberia 70% of the population is reported to
consume some bushmeat or to sell it. Estimates of game
meat consumption in a variety of other African countries
are given in Table 26.2. In Nicaragua wildlife provided
over 98% of the meat and fish consumed by the Miskito
Indians (Nietschmann, 1973). Studies of hunting and wild
meat consumption in Sarawak have estimated the total value
of wild meat production of about 18,000 tonnes as having
a replacement value (cost of domestic substitutes)
approaching M$100 million. Even in an industrialised
country such as the USA, sport hunting of large ungulates
alone was estimated to yield 150,000 tonnes of meat a year
with a replacement value of US$450 million (Payne, 1989).
In Sweden, the shooting by sport hunters of 186,000 Elk
Alces alces in 1983 yielded 3.0-3.4kg of meat of this single
species per head of population (Bubenik, 1989). Many
different species of wild animal are exploited as sources of
food, providing a variety of dietary essentials such as
protein, fats, and oils. The most conspicuous terrestrial
source of wild animal protein is medium to large mammals.
Subsistence hunters generally take more mammals than
birds, and more birds than reptiles. This is reflected in the
number of species taken: the bush people of Suriname take
at least 27 mammal species, 24 birds, three turtles and two
species of lizards (Redford and Robinson, 1991).
Amongst mammals, ungulates, primates, and large rodents
all figure prominently in the bushmeat trade in Africa (see
Table 26.3) and South America. Edentates (anteaters and
armadillos) are also taken in the New World, while fruit
bats of the genus Pteropus are considered a delicacy in
Oceania.
Birds generally provide meat and eggs for human
consumption, but even their nests may be eaten in the case
of the cave swiftlets of Southeast Asia. For example, the 49
species of Cracidae constitute an important source of meat
for the campeseino and indigenous Indian populations. The
eggs of the Greater Rhea Rhea americana, Black-bellied
Whistling-duck Dendrocygna autumnalis, and flamingos are
all collected for consumption in South America (Redford
and Robinson, 1991). The eggs and young of seabirds are
eaten in many parts of the world, probably because the
colonial nesting habit makes them particularly easy to
collect in large quantities.
Among reptiles, monitor lizards are widely eaten in Africa,
iguanas in South America, and sea turtles, particularly
Green Turtles Chelonia mydas, provide meat for many
littoral peoples worldwide. In many areas, reptiles are an
even more important source of eggs than birds: eggs of sea
turtles, freshwater turtles, Green Iguana Jguana iguana and
the tegu Tupinambis spp. are all consumed by local people
and exploited as a source of income.
2. Uses and Values of Biodiversity
Table 26.1 Daily per capita consumption of animal protein (g) in countries obtaining
more than half of their average supply from wild animals
COUNTRY TOTAL ANIMAL GAME MEAT FISH AND TOTAL WILD WILD AS %
PRODUCTS SEAFOOD MEAT OF TOTAL
ASIA
Bangladesh 6.7 - 3.5 3.5 52.2
Indonesia 5.3 - 3.6 3.6 67.9
Korea, DPR 12.1 - 8.0 8.0 66.1
Korea, Rep 13.1 - 9.0 9.0 68.7
Malaysia (Sabah) 22.1 - 11.3 11.3 Bilist
Malaysia (Sarawak) 15.5 - 9.3 9.3 60.0
Maldives 30.6 - 28.4 28.4 92.8
Philippines 16.9 - 8.9 8.9 52a,
Thailand 13.2 - 6.8 6.8 51.5
Viet Nam 14.0 - 7.8 7.8 55.7
OCEANIA
Papua New Guinea 18.2 2.6 8.3 10.9 59.9
AFRICA
Benin 8.3 161 3.7 4.8 57.8
Congo 11.8 2.0 6.9 8.9 75.4
Ghana 15.2 1.4 10.0 11.4 75.0
Liberia 9.2 1.4 4.6 6.0 65.2
Senegal i7éaal - 9.6 9.6 56.1
Sierra Leone 10.0 0.3 phe? TSI 75.0
Togo 6.8 0.9 3.1 4.0 58.8
Zaire 7.1 8) 2.6 4.5 63.4
Source: Modified from Prescott-Allen, R. and Prescott-Allen, C. 1982. What’s Wildlife Worth? Earthscan; FAO, 1977. Provisional Food Balance
Sheets: 1972-74 average.
Table 26.2 Estimated annual game output and per capita supply in selected African
countries
OUTPUT SUPPLY CONTRIBUTION OF GAME
(thousand PER CAPITA MEAT TO PER CAPITA
tonnes) (kg) ANIMAL PROTEIN SUPPLY
1972-74 1977 1972-74 1977 (%)
Angola 6 6 1.0 0.9 4.5
Benin 6 6 2.2 1:9 13.2
Botswana 5 6 7.6 7.5 15.8
Cameroon 4 4 0.6 0.6 2.8
Chad 3 3 0.8 0.7 Kh)
Congo 5 6 4.0 4.0 16.9
Céte d'Ivoire 13 13 2.8 2.5 7.4
Ethiopia 7 7 0.2 0.2 1.0
Gambia 1 1 1.7 1.8 7.8
Ghana 26 28 2.8 7a) 9.2
Guinea 4 4 0.9 0.8 10.0
Kenya 7 a 0.6 0.5 2.6
Lesotho 3 4 3.0 2.9 13.4
Liberia 5 5 2.8 3.0 shes
Namibia 2 2 2.6 2.6 3.9
Nigeria 87 95 1.2 1.2 13.0
Rwanda 5 6 1.3 1.3 25.0
Sudan 6 7 0.4 0.3 1.0
Tanzania 7/ 8 0.5 0.5 1.6
Togo 4 4 1.9 1.7 13.2
Uganda 12 14 al! 1.2 5.0
Zaire 90 68 3.9 2.6 26.8
Zambia 17 20 iy! Sr7, 13.4
Source: Modified from Prescott-Allen, R. and Prescott-Allen, C. 1982. What’s Wildlife Worth? Earthscan; FAO data.
The only amphibians widely used as a source of food are export to supply frogs’ legs to the gourmet market in
frogs. The frogs’ leg trade, mainly based on wild-caught Europe and North America, although some local
individuals of the genus Rana, is economically important in consumption does occur.
Asia. The majority of the specimens caught are destined for
360
Animal Use
Table 26.3 Wild animals of the bushmeat trade in Ghana*
NORTHERN GUINEA
SEMI-DECIDUOUS
COASTAL PLAINS
SAVANNA FOREST SAVANNA
Warthog Baboon Grasscutter
Baboon Warthog Giant Rat
Hartebeest Grasscutter Royal Antelope
Bushbuck Hartebeest Bushbuck
Crowned Duiker Kob Bat
Aardvark Bushbuck Green Monkey
Grasscutter Roan Antelope Crowned Duiker
Roan Antelope Aardvark Black Duiker
Buffalo Waterbuck Red River Hog
Waterbuck Oribi Monitor Lizard
Source: Sale, J.B. 1981. The Importance and Values of Wild Plants and Animals in Africa. Part 1. IUCN, Gland, Switzerland.
Note: * Top ten species, listed in descending order of importance
Invertebrates
The molluscs and the arthropods include many species used
as a food resource. Marine and freshwater molluscs are
more important to human nutrition on a global basis than
terrestrial species, but in certain areas the latter may figure
prominently in the diet. Thus the Giant African Land Snail,
of the genus Achatina, is eaten on a large scale in West
Africa and is immensely popular with people in central
Ghana and parts of Nigeria. It has a protein value nearly
equivalent to beef. Non-insect arthropods used for food
include land crabs, centipedes, woodlice and large spiders.
Insects are an important supplementary source of calories
and protein in many regions of the world. Examples of
some of the 500 or so species known to be consumed are
given in Table 26.4 with an indication of the region where
they are eaten.
Insects of most major orders are eaten, but the most widely
used species are those, such as termites, which habitually
occur in very large numbers in one place, or which
periodically swarm, such as locusts, or large species such
as saturniid moth larvae. The seasonal abundance of certain
species makes them especially important at times of year
when other food resources may be lacking.
Orthoptera (grasshoppers, crickets etc.) are a valued food
for many peoples. Swarming locusts can easily be gathered
by the sackful, are easily dried for storage and can be a
valuable resource to help tide over hard times.
The Lepidoptera (butterflies, moths) is probably the order
containing the largest number of species eaten. Especially
popular in Asia and Africa are many species of saturniid
moths which have large fleshy larvae. Some species are
dried and sold to quite a large market and are important in
the local economy. The pupae of various species of silk
moths are consumed in much of Southeast Asia, partly as
a by-product of the silk industry. The pupae are killed by
immersing them briefly in hot water before the silk is
wound off, after which they may be eaten and thus provide
361
an important source of nutrients for many silk workers
(Taylor, 1975; Vane-Wright, 1991).
Isoptera (termites) are eaten almost everywhere they occur.
In some areas only the swarming reproductive termites are
taken; in others, the nests are dug out and all stages are
eaten (TFIN 5).
The Hymenoptera (bees, wasps and ants) is another widely
utilised order. Honey produced by bees from nectar and
pollen is prized, and is one of the most widely accepted
insect products (Table 26.5). However, the honey can be
less important than the bee brood which is collected with
the honey. These immature stages in the comb are eaten by
many indigenous peoples and are highly nutritious (TFIN
3). Similarly, wasp brood is eaten in some areas, and less
commonly, adult wasps. Larger ants are quite common as
food items. In Australia, honeypot ants are or were popular
with Aboriginal peoples and leafcutter ants are commonly
eaten in parts of the Americas.
Beetles (Coleoptera) are eaten both as larvae and adults. In
Thailand, several species are eaten as adults and can often
be bought live in markets (Watanabe and Satrawaha, 1984).
The abdomens of adult rhinoceros beetles are also eaten by
the Yukpa of Venezuela and Colombia (Ruddle, 1973).
Despite the widespread use of insects and other
invertebrates for food, they represent an under-exploited
resource. Great fecundity is a feature of insects giving
potential for wild harvesting and farming. There have been
several studies on large-scale insect production both as
human food and for animal feed (Maitipe, 1984; TFIN 1
and 4). Many species which are agricultural pests are also
used as a food resource in some part of their range, or have
the potential to be utilised. Palm grubs (weevil larvae),
which are a pest of coconuts and oil palms throughout the
tropics but are considered a delicacy by indigenous peoples,
are a case in point. Several Indian tribes in the American
tropics ‘farm’ the larvae in that logs are prepared as laying
sites and then mature larvae are collected several months
later. Several hundred grammes of larvae can be collected
per log (TFIN 2).
2. Uses and Values of Biodiversity
Table 26.4 Selected insects used as a human food source
ORDER
FAMILY
Odonata (Dragonflies)
Aeschnidae
Libellulidae
(general Odonata)
Blattaria (Cockroaches)
Blattidae
Mantodea (Mantids)
Mantidae
lsoptera (Termites)
Rhinotermitidae
Termitidae
Orthoptera (Grasshoppers, crickets etc)
Tettigoniidae
Gryllidae
Acrididae
Phasmoptera (Stick and leaf insects)
Phylliidae
Phasmatidae
Hemiptera (Bugs)
Belistomatidae
Corixidae
Pentatomidae
Homoptera (Bugs)
Cicadidae (cicadas)
Neuroptera (Lacewings etc)
Corydalidae
Coleoptera (Beetles)
Dytiscidae
Scarabaeidae
Buprestidae
Tenebrionidae
Bruchidae
Curculionidae
Stratiomyidae
Trichoptera (Caddis flies)
Hydropsychidae
Lepidoptera (Butterflies and moths)
Hepialidae
Cossidae
Hesperidae
Megathymidae
Pieridae
Bombycidae
Saturniidae
SCIENTIFIC NAME
Blatta orientalis
Periplaneta
Hierodula sternosticta
Mantis religiosa
Coptotermes formosanus
Macrotermes
Termes flavicolle
Conocephalus angustifrons
Acheta
Gryllotalpa africana
Aidemonaazteca
Locusta
Oryxa
Tropidacris l/atreillei
Haaniella grayi
Eurycantha horrida
Platycrana viridana
Lethocerus indicus
Corixa femorata
Erthesina fullo
Corydalus
Cybis
Cybista hova
Copris
Heliocopris bucephalus
Oryctes
Podischnus agenor
Sternocera
Tenebrio
Caryobruchus
Anthonomus
Phyncophorus
Chrysochlorina
Leptonema
Hepialus amoricanus
Xyleutes leuchomochla
Acentrocneme hesperialis
Aegiale hesperialis
Eucheira socialis
Bombyx mori
Athletes semialba
Bunaeopsis aurantiaca
Cirina forda
Gonimbrasia
Gynasia maja
Lobobunaea saturnus
Urota sinope
362
LIFE STAGE
NORMALLY
CONSUMED
>
>
=
lt -rprprrrprrp
Ce ed A eit eee eh Tl eal ea dD od
AREA WHERE IT IS EATEN
Southeast Asia
Southeast Asia
West Africa, Asia, New Guinea
Southeast Asia, Madagascar
Southeast Asia
Asia, Australia
New Guinea
Southern Africa
Asia
Africa
South America
Tropical South America
Southeast Asia
Southeast Asia
Tropical South America
Asia
Asia
Tropical South America
Southeast Asia
New Guinea
Southeast Asia
Asia, Southeast Asia
Central America
Asia
New Guinea
Tropical South America
Southeast Asia
Madagascar
Southeast Asia
Asia, Southeast Asia
Africa, Asia and western Pacific
Tropical South America
Southeast Asia
South-west Asia
Tropical South America
Tropical South America
Tropics worldwide
Tropical South America
Tropical South America
Asia
Australia
Central America
Central America
Asia
Central Africa
Central Africa
Central Africa
Africa
Central Africa
Central Africa
Central Africa
Animal Use
Table 26.4 Selected insects used as a human food source (continued)
Lepidoptera (Butterflies and moths) (continued)
Notodontidae Antheua insignata L Central Africa
Elaphrodes lactea L Central Africa
Thaumatopoeidae Anaphe panda L Central Africa
Noctuidae Laphygma frugiperda L Tropical South America
Nyodes prasinodes L Central Africa
Hymenoptera (Ants, wasps, bees etc)
Formicidae Atta A Tropical South America
Liometopum apiculatum A?,E,L?,P? Central America
Melophorus bagoti A Australia
Oecophylla smaragdina EF Southeast Asia
Vespidae Mischocyttarus L Tropical South America
Polistes E Tropical South America
Polybia ignobilis L Tropical South America
Vespula lewisi A,L,P Asia
Apidae Apis dorsata L,P Southeast Asia
Apis laboriosa L,P Asia
Trigona A?,L Tropical South America, Australia
Source: See references; drawn from various texts on entomology.
Notes: A = adult, E = egg, L = larva, N = nymph, P = pupa.
Table 26.5
1989
ASIA
China 177,000 *
India 50,000 F
Turkey 40,000 F
USSR 230,000
EUROPE
Czechoslovakia 10,000 F
France 26,000 F
Germany 23,300 F
Greece 11,650 F
Hungary 16,000 F
Poland 15,000 F
Romania 17,000 F
Spain 21,000 F
Countries with honey production at or in excess of 10,000 tonnes in
NORTH AND CENTRAL AMERICA
Canada 28,100
Mexico 52,530 *
United States 80,000
SOUTH AMERICA
Argentina 38,000 *
Brazil 16,000 F
OCEANIA
Australia 22,619
AFRICA
Angola 15,000 F
Egypt 13,000 F
Ethiopia 22,600 F
Kenya 16,000 F
Tanzania 14,000 F
Source: FAO, 1990a. FAO Yearbook: Production 1989. Food and Agriculture Organization of the United Nations, Rome.
Notes: F = FAO estimate; * = unofficial figure.
Nutritionally, as demonstrated in Table 26.6, insects
compare well with other animal products; furthermore, the
efficiency of food conversion to biomass is also favourable
(Table 26.7).
Trade in meat products
In all its nutritional forms, wildlife has great economic
value to local people. Thus Ajayi (1971) calculated that the
annual replacement value of wild animal protein (inclusive
of bushmeat, wildfowl and fish) used in Nigeria totalled
some £30 million which at the time was approximately
equal to 4% of Nigeria’s gross domestic product.
Many species are also traded in local markets, providing
direct revenue for hunters and traders alike. Around
Kisangani (Zaire), the village hunters take the higher value
363
carcasses, especially duikers, to the towns for sale, while
the rodents and other smaller animals are mostly consumed
by the hunters and their families (Colyn et al., 1988). The
advent of efficient transportation and storage has allowed
the development of commercial food industries based on
international trade in wildlife meat. Green Turtles Chelonia
mydas have long been exported from the Caribbean and
Indian Ocean to Europe and elsewhere for the gourmet
trade but this is now prohibited by the Convention on
International Trade in Endangered Species (CITES). Most
species traded for meat are relatively common and therefore
not listed in the CITES statistics. However Customs
statistics often contain a category for "game meat", and an
analysis of this produced an estimate of an annual average
trade of some 32,000 tonnes (Table 26.8). The main
exporting countries were Argentina, the UK and several
Eastern European countries, particularly Hungary and
2. Uses and Values of Biodiversity
Table 26.6 Nutritional values of selected vertebrate and invertebrate products
LIFESTAGE PROTEIN FAT CARBO- WATER ASH CALORIES
(%) (%) HYDRATE (% weight) (minerals) 1100g
(%) (%)
Vertebrate products
Beef 17.4-19.4 15.8-25.1 0.0 56.7-63.9 0.8-0.9 225-301
Lamb 15.4-16.8 19.4-27.1 0.0 56.3-62.5 1.2-1.3 247-310
Pork 14.6-16.7 22.7-31.4 0.0 52.6-59.5 1.0-1.2 276-346
Chicken 20.6-23.4 1.9-4.7 0.0 73.7 1.0 117-130
Fish
Lake trout 18.3 10.0 0.0 70.6 ua 168
Halibut 20.9 1.2 0.0 76.5 1.4 100
Milk 3.5 3.7 4.9 87.2 0.7 66
Eggs 12.9 11.5 0.9 73.7 1.0 163
Invertebrates
lsoptera
(Living, species not known) A 23.2 28.3 44.5 347
(Fried, species not known) A 36.0 44.4 6.0 561
Orthoptera
(Living: mixture of species) A 15.3-46.1 2.4-9.6 6.8-7.5 10.5-70.6 0.8-5.0
(Sun-dried: mixture of A 49.7-75 10.1-18.4 6.4-16.1 5.0 3.7-18.9
Coleoptera F,
Scarabaeidae
Lachnosterna sp. it 11.1 3.1 2.3 79.9 2.0
A 20.1 4. 0.3 69.4 1.6
Curculionidae
Polycleis equestris A 30.3 eae. 51.8
Diptera
Muscidae
Musca domestica P 63.1 15.5 3.9 5.3
Lepidoptera
Bombycidae
Bombyx mori P 23.1 14.2 60.7 AS 207
Saturnidae
Athletes semialba L 15.5 4.5 1.2 78.0 0.8 504"
Bunaea alcinoe L 5.9 0.9 1.7 91.0 0.4 443"
Gonimbrasia richelmanni L 15.9 ye | 0.7 80.0 1.3 447*
Gynanisa maja L 10.1 3:3 1.0 84.5 1.1 495°
Imbrasia rubra L 11.8 72 | 0.2 83.0 1.3 445"
Notodontidae
Drapedites uniformis (L 10.8 4.0 331 79.5 1.1 452"
Elaphrodes lactea L 16.3 5.9 1.8 72.0 1.2 461"
Hymenoptera
Apidae
Apis mellifera L 15.4 3.7 0.4 77.0 3.0
P 18.2 2.4 0.8 70.2 2.2
Araneae
Theraphosidae
Melopoeus albostriatus A 63.4 9.8
Source: Adapted from Malaisse and Parent, 1980. Les chenilles comestibles du Shaba meridional (Zaire). Naturalistes Belges 61(1):2-24 and Taylor,
R.H. 1975. Butterflies in my Stomach. Woodbridge Press Publishing Co., Santa Barbara, California (see also table in The Food Insects Newsletter,
IV (1), on fatty acids). Notes: * Dry weight.
Table 26.7 Efficiency of food conversion for selected animals
Chicken (broilers)
Turkeys
Sheep and lambs
Beef cattle and calves
Pigs
Fish and shellfish
Blattaria (Cockroaches)
Blatella germanica
Orthoptera (Grasshoppers etc.)
Gryllus domesticus
Melanoplus bilituratus
Schistocerca gregaria
Phasmoptera (Stick Insects etc.)
Carausius morosus
Hemiptera (True Bugs)
Cimex lectularius
Phonoctonus nigrofasciatus
Rhodnius prolixus
Stalia major
Coleoptera (Beetles)
Lasioderma serricorne
Efficiency %*
38-40
21
5.3
21-41
33-53
19-24
41-54
21.4
Paropsis atomaria
Tribolium confusum
Diptera (Flies)
Cochliomyia hominivorax
Lepidoptera (Moths etc.)
Aglais urticae
Agrotis orthogonia
Bombyx mori
Chilo suppressalis
Dendrolimus pini
Galleria mellonella
Hepialus humuli
Hyphantria cunea
Malacosoma neustria
Mamestra brassicae
Phalera bucephala
Pieris brassicae
Prodenia eridania
Protoparce secta
Smerinthus populi
Tineola bisselliella
Efficiency %*
Source: Compiled from data in Taylor, R.H. 1975. Butterflies in my Stomach. Woodbridge Press Publishing Co., Santa Barbara, California.
Note: * % of food consumed converted into animal tissue (optimum).
364
Table 26.8
NET EXPORTERS 1980 1981
Algeria 2 -
Argentina 12,098 10,468
Australia 382 1,860
Austria 1,015 999
Brazil = =
Bulgaria 108 58
Canada 24 --
Chile - 50
China 1,444 876
Czechoslovakia 977 918
Finland 24 53
Germany, DR -- -
Greece oa =
Greenland 39 51
Hungary 3,086 2,534
Iceland - -
Ireland 26 26
Israel - 8
Mongolia 121 133
Morocco -- -
Netherlands * 627
New Zealand 1,017 1,641
Norway x
Poland 1,729 2,217
Romania 678 429
South Africa 3,479 2,197
Spain 1,296 1,560
Sweden 270
Tunisia = 26
Turkey 33 --
UK 5,041 4,761
Uruguay 358 237
USA 37 37
USSR 883 1,133
Yugoslavia 1,081 1,144
Country unknown 44 =
Total tonnes 35,292 34,043
Animal Use
Estimated minimum net exports of game meat (tonnes)
1982 1983 1984 1985
9,304 9,599 8,986 11,627
1,403 337 146 103
873 953 192 718
10 22 20 31
31 150 194 174
49 189 137 147
853 1,719 2,110 1,528
1,180 1,184 1,488 1,316
21 31 95 59
= = = 32
= = 30 17
51 41 56 38
2,369 2,934 2,763 3,097
= = = 16
52 70 54 63
138 167 = 166
be =: 14 15
746 710 350 313
1,197 929 831 1,291
* * 6 *
1,858 3,449 3,119 2,850
523 291 293 89
1,546 560 835 1,033
1,620 1,574 1,758 1,632
= : 5 :
< 28 34 10
3,857 4,061 4,788 4,695
181 218 280 495
— * . *
1,321 578 1,075 1,433
918 850 1,040 1,278
" 15 43 56
30,101 30,659 30,737 34,322
Source: Luxmoore, R.A. 1989. International trade. In: Hudson, R.J., Drew, K.R. and Baskin, L.M. (Eds), Wildlife Production Systems: economic
utilisation of wild ungulates. Cambridge University Press, Cambridge, UK.
Note: * Net importers in this year.
Poland. Exports from Argentina comprise largely Cape
Hare Lepus capensis, and from the UK, Red Deer Cervus
elaphus and a variety of game birds. Because of veterinary
health controls, international trade in fresh meat is very
much more closely controlled than most animal products.
This effectively prohibits exports of game meat from much
of Africa to Europe, which constitutes the main market, and
explains why South Africa is virtually the only exporter
listed in Table 26.8.
FOOD: FISHERIES
Global fish production exceeds that of cattle, sheep, poultry
or eggs, and is the largest source of either wild or domestic
animal protein for the world’s expanding human population
(Norse, 1992). It is particularly important in the developing
countries, as is evidenced by the large contributions fish
and seafood make to the totals in Table 26.1.
The fisheries industry is a large and expanding one. Annual
world landings of aquatic resources have increased more
than four-fold in the last 40 years, from 21.9 million tonnes
per year between 1948 and 1952 to 99.5 million tonnes in
1989 (FAO, 1990b, 1991a). The majority of these landings
365
originated from marine fisheries and were destined for
human consumption. Marine landings comprised 85.8
million tonnes (86.2% of total landings in 1989) while
inland fisheries (aquaculture and capture fisheries)
accounted for the remaining 13.8 million tonnes, or 13.8%
(FAO, 199ia). Almost 70% (69.2 million tonnes) of total
landings were used for human consumption, while the
remainder were used for animal feed, fertilizer etc. The
vast majority of the catch (92.1%) comprises fishes
(marine, diadromous and freshwater), with molluscs,
crustacea and other animals being relatively unimportant in
terms of global landings (Fig. 26.1). However, many of
these groups command high prices per kg and have a
disproportionately high economic value (FAO, 1991b).
Distribution of marine fisheries
The location of the world’s marine fisheries is governed
principally by the distribution of the floating plants on
which they depend for food. Phytoplankton production is
principally dependent on adequate supplies of nutrients, and
is largest in areas of upwelling.
Climatic fluctuations can greatly alter the pattern of ocean
2. Uses and Values of Biodiversity
Figure 26.1 World nominal catches in
1989 by groups of species
Crustacea (3.0%)
Freshwater fishes (11.5%)
Miscellaneous (0.3%)
Diadromous fishes (7.9%)
UY
Molluscs (4.5%)
Marine fishes (72.7%)
Source: FAO 1991a. FAO Fishery Statistics Yearbook: catches and
landings 1989. Vol. 68. FAO, Rome.
Note: Total world nominal catch = 99,534,584 tonnes.
circulation, and hence fisheries production, around the
world. Perhaps the most famous of these events is the
disruption in some years of the circulation pattern off the
coast of Peru, a phenomenon known as ‘El Nifio’, which
intermittently leads to the near total collapse of the coastal
fisheries.
The relative importance of the catches in the different
fishing areas reflect the differences in production. The four
major fishing areas (as defined for statistical purposes by
FAO) in descending order of annual tonnage of landings are
the North-west Pacific, the South-east Pacific, the
North-east Atlantic and the Western Central Pacific (Fig.
26.2, Table 26.9).
Composition of marine fisheries
Although there are approximately 22,000 species of fish, of
which more than 13,000 are marine (Nelson, 1984), only a
very small fraction are of major commercial importance.
FAO statistics (FAO, 1991a) break down aquatic animals
and plants into 980 "species items" (species, genera, or
families) which are then further categorised into 51 groups
of species. Of these, only 17 contributed more than 1% (=
one million tonnes) towards total recorded world landings,
which approached 100 million tonnes in 1989 (Fig. 26.3).
The most important groups were the herrings, sardines and
anchovies, of which 24.5 million tonnes were landed in
1989, followed by the cods, hakes and haddocks, of which
12.8 million tonnes were landed.
The fisheries industry is based on a remarkably small
number of species. Over one million tonnes each of 12
individual fish species (10 marine and two freshwater, see
Table 26.10) were caught in 1989: together these comprised
34.7 million tonnes, or 34.9% of the total world catch. The
single largest species fishery was the Alaska Pollock
Theragra chalcogramma of which 6.3 million tonnes were
landed, while over five million tonnes of both the
Anchoveta Engraulis ringens and Japanese Pilchard
Sardinops melanostictus were also caught. Of the six largest
fisheries, five are located in the Pacific (three in the South-
east Pacific and two in the North Pacific) while one is from
the North Atlantic.
366
Recent trends in marine fisheries
Reported world landings have generally increased over the
past 25 years. During the 1960s (Fig. 26.4), total landings
increased steadily as new stocks were discovered, while
improved fishing technology and an expansion of fishing
effort enabled fuller exploitation of existing stocks of both
pelagic (surface water or open sea) and demersal (deep
water or bottom-dwelling) species. Long-range fleets
increased in size during this period, concentrating their
efforts in the richest ocean areas, and were largely
responsible for the rapid increase in world catches.
In the 1970s, following the collapse of the Peruvian
anchovy fishery there was very little increase in the total
catch. Landings of most demersal fish stocks remained
relatively constant, implying that they were close to full
exploitation and, whilst landings of pelagic fish stocks
changed from one species to another in certain areas, there
was no appreciable change in total pelagic landings (FAO,
1990b). Long-range fleets continued to expand in
importance.
The 1980s once again saw a period of continuous growth
(averaging 3.8% a year) in world landings. Because most
demersal stocks were (and still are) fully fished, shoaling
pelagic species provided most of the increase in fish
production. In fact, just three pelagic species (Peruvian
Anchovy Engraulis ringens, South American Sardine
Sardinops sagax, and Japanese Sardine Sardinops
melanostictus) and one semi-demersal species (Alaska
Pollock Theragra chalcogramma) accounted for 50% of the
increase in world landings during the 1980s (FAO, 1990b).
Most of this increase appears to have been because of
favourable climatic effects on stock sizes rather than new
fishery developments or improved management practices
(FAO, 1990b).
A concurrent change in the fishing industry in the 1980s
was the increase in levels of national and international
controls designed to ensure the conservation of fish stocks.
This reduced the importance of long-range fishing in many
areas and allowed the development of short- or medium-
range fishing fleets, (FAO, 1990b). Thus in the early 1970s
long-range catches formed 79 % of the North-eastern Pacific
catch, but had declined to only 8% in 1988, having been
replaced by local fleets and joint fishing ventures (FAO,
1990b).
The regional trend in marine landings over the period 1983-
1989 was upwards in all but three of the FAO designated
fishing areas: the Mediterranean and Black Sea, where
output was relatively stable; and the North-east Atlantic and
Western Central Atlantic where output declined slightly
(Fig. 26.2, Table 26.9). The largest increases occurred in
the North-east Pacific and the South-east Pacific.
Although annual world landings of aquatic resources have
grown steadily since the 1950s (Fig. 26.4) fishery resources
around the world are now thought to be close to their
maximum catch limits, and many show signs of biological
degradation (FAO, 1990b). Total world marine catch in
1989 was 85.8 million tonnes (FAO, 1990b) and it has been
Animal Use
Figure 26.2 Catches in FAO fishery areas, 1984-1989
v
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DDABBABAALLA
:
,
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367
2. Uses and Values of Biodiversity
Table 26.9 World nominal catches of fish, crustaceans and molluscs, 1984-1989
FAO FISHING AREA YEAR
Region No. 1984 1985 1986 1987 1988 1989
INLAND
Africa 1 1535000 1542900 1667200 1746400 1842800 1871400
America, North 2 433900 443300 484500 572900 535100 528800
America, South 3 338500 328900 362100 386200 356400 321900
Asia 4 6405800 7025400 7840400 8564000 9175900 9535700
Europe 5 413800 430600 459400 448700 474800 476000
Oceania 6 20000 20400 20500 22300 23400 23100
USSR 7 881500 905600 926900 988400 995600 1019700
MARINE
Arctic Sea 18 0 0 0 te) te) t)
Atlantic, Northwest 21 2734300 2870000 2961900 3079800 3020900 3079300
Atlantic, Northeast 27 11460600 11118700 10589100 10457400 10567500 9931000
Atlantic, Western Cental 31 2598800 2246400 2044800 2144000 1874500 1791900
Atlantic, Eastern Central 34 2692000 2844800 3027100 3194100 3533500 3702300
Mediterranean & Black Sea 37 2016000 1979000 2012400 1946500 2071300 1673000
Atlantic, Southwest 41 1567200 1700700 1846500 2370300 2329900 2254200
Atlantic, Southeast 47 2143700 2104100 2125100 2728800 2499400 2095000
Atlantic, Antarctic 48 225200 228200 462000 434400 443100 465200
Indian Ocean, Western 51 2542900 2654200 2661600 2719600 2985300 3290900
Indian Ocean, Eastern 57 2328000 2271300 2513900 2656600 2720400 2758200
Indian Ocean, Antarctic 58 35600 31300 37200 39100 14900 31400
Pacific, Northwest 61 23717100 23841700 25709600 25848300 26658000 26310500
Pacific, Northeast 67 2689200 2882100 3204900 3447400 3338600 3290700
Pacific, Western Central 71 5867500 5903500 6416900 6829000 6990800 7076800
Pacific, Eastern Central a, 1213600 1700700 1642100 1753000 1655400 1705200
Pacific, Southwest 81 612800 577500 755100 907900 969000 990600
Pacific, Southeast 87 9465800 10741600 13000500 10988000 13665800 15310600
Pacific, Antarctic 88 800 4700 3900 400 0 1100
Total 73911100 75700500 81014600 81544600 85338300 85757900
Source: FAO 1991a. FAO Yearbook, fishery statistics (catches and landings). Vol. 68, 1989. FAO, Rome.
Figure 26.3 FAO species groups contributing over 1% to world catches in 1989
25
w
to}
ae Marine fishes
Freshwater fishes
Molluscs
(ee) Crustaceans
le ro Diadromous fishes
fe) é ;
=
oa
World catch (million tonnes)
ra)
8 2 4 8 £ 8 3 r 3 8 & 2 2 3 8 8 £
a ee oe ee ae ee
eB = ry ts as tS a gS 5 5 ~ Boy
ra] gz © ® S . = = 2 i] a 2 S o > o
a ae eS ae lee ee Re Se = ae
= 5 2 3 s Pee 2 a 5 a 2 3
3 = © = 2 ° . ° © =
3 E = 8 E LS rs) o — eg 3 a
= = a 7 3 = S © < = z 2
= o = E ry 2 ¢ £ = ~ 9
5 £ 3 = a £ ae S 2 ry a o ° ~
a = 9 § c Ee 2 © i= c ° £ no
eee Ls A re I Se a 2 ¢ 3
c ° Ss Fy ro) 2 G a g £ e
BS 9 Oo Sys ee et Se ae > 2 s 6§ 3
c Oo 7. c 2 a a n = o
1: 2 2 = 6 F © ry o ira
6 = « © a o =|
=x 4 = * 5
2 B 8 oc
= 5 s 7)
Species group
368
Animal Use
Table 26.10 World total and area nominal catches for the principal fishery species*
SPECIES
Alaska Pollock
Anchoveta
Japanese Pilchard
South American Pilchard
Chilean Jack Mackerel
Atlantic Cod
Chub Mackerel
Atlantic Herring
European Pilchard
Silver Carp
Skipjack Tuna
Common Carp
Source: FAO 1991a. FAO Yearbook Fishery Statistics: catches and landings 1989. Vol. 68. FAO, Rome.
Theragra chalcogramma
Engraulis ringens
Sardinops melanostictus
Sardinops sagax
Trachurus murphyi
Gadus morhua
Scomber japonicus
Clupea harengus
Sardina pilchardus
Hypophthalmichthys molitrix
Katsuwonus pelamis
Cyprinus carpio
AREA
Pacific, Northwest
Pacific, Northeast
Pacific, Southeast
Pacific, Northwest
Pacific, Northeast
Pacific, Southeast
Pacific, Southeast
Atlantic, Northwest
Atlantic, Northeast
Atlantic, Northeast
Atlantic, Western Central
Atlantic, Eastern Central
Mediterranean and Black Sea
Atlantic, Southwest
Atlantic, Southeast
Indian Ocean, Western
Pacific, Northwest
Pacific, Northeast
Pacific, Western Central
Pacific, Eastern Central
Pacific, Southwest
Pacific, Southeast
Atlantic, Northwest
Atlantic, Northeast
Atlantic, Northeast
Atlantic, Eastern Central
Mediterranean and Black Sea
America, North (inland)
Asia (inland)
Europe (inland)
Atlantic, Northeast
Atlantic, Northwest
Atlantic, Northeast
Atlantic, Western Central
Atlantic, Eastern Central
Atlantic, Southwest
Atlantic, Southeast
Indian Ocean, Western
Indian Ocean, Eastern
Pacific, Northwest
Pacific, Northeast
Pacific, Western Central
Pacific, Eastern Central
Pacific, Southwest
Pacific, Southeast
Africa (inland)
America, North (inland)
America, South (inland)
Asia (inland)
Europe (inland)
USSR (inland)
Atlantic, Northeast
Mediterranean and Black Sea
Note: * Those with catches of over one million tonnes; data for 1989.
369
AREA CATCH WORLD CATCH
(tonnes)
4,741,659
1,517,399
5,407,527
5,111,525
58
4,196,169
3,654,628
630,170
1,152,412
6,889
607
313,727
28,095
13,150
30,200
98
986,333
1,470
697
43,724
973
245,107
275,110
1,337,076
172,842
973,643
253,545
1,509
1,339,556
18,650
9
41
5,931
4,203
84,851
23,053
585
217,323
14,074
120,135
682
589,247
78,639
10,461
30,896
812
25,050
688
679,315
123,885
255,706
70
15
(tonnes)
6,259,058
5,407,527
5,111,583
4,196,169
3,654,628
1,782,582
1,671,070
1,612,186
1,400,030
1,359,724
1,180,121
1,085,341
2. Uses and Values of Biodiversity
estimated that there is now little scope for increased catches
of any of the traditionally fished marine species. There is an
increasing need for conservation measures to protect and
manage fish stocks in order to sustain current levels of take
and rehabilitate degraded fisheries (FAO, 1990b).
The most important step to facilitate the sustainable
exploitation of fish stocks has been the establishment by
coastal states of jurisdiction up to 200 miles from their
shores; 99% of the marine fisheries catch is currently taken
within this limit (FAO, 1990b). Most countries are now
declaring or have declared 200-mile fishing exclusion zones
around their coasts, providing increased potential for
tational and sustained use of resources, (FAO, 1981).
Further discussion of fisheries management practices, with
particular reference to international agreements, is provided
in Part 3.
Trends in fish stocks
All species of fish are subject to population and recruitment
fluctuations which vary according to the species’ biology,
migratory habits, food resource availability, natural
hydrographic factors, fishing practices and management.
Brief details of two stocks are given below in order to
illustrate the nature of such fluctuations.
Atlantic Herring (Clupea harengus harengus)
The Atlantic Herring provides an example of an over-
exploited fish population that has recovered under sound
management. It is widely distributed on both sides of the
North Atlantic in many reproductively independent groups
(races). Following a long period of overfishing and the
failure of management controls, most stocks declined to
very low levels in the 1970s. At that point, bans and
subsequent catch quotas were introduced which allowed
stocks to recover (Fig. 26.5).
The North Sea Herring reached a minimum of 75,000
tonnes in 1975, but a ban on fishing from 1977 to 1981 has
allowed the population to build up to 1.4 million tonnes
which, although lower than the post-War size, is about the
same level as in the 1950s and 1960s prior to the collapse
(Corten and van de Kamp, 1991).
The Norwegian spring-spawning stock, once the largest
herring stock in the world with a spawning biomass of 10
million tonnes in 1957, collapsed to virtually zero in the
1970s as a result of excess fishing pressure and subsequent
poor recruitment. Landings decreased to only
10,000-20,000 tonnes between 1972 and 1983, but
management measures (e.g. fishing quotas, minimum mesh
sizes etc.) permitted the stock to recover. Landings in 1988
were recorded as 125,000 tonnes and the spawning stock
biomass estimated at 1.3 million tonnes (ACFM, 1991).
Certain races have never recovered from the earlier
overfishing: the spring-spawning stock of Icelandic Herring
is now effectively extinct, but Jakobsson (1985) believes
that the failure of this stock to recover may have been
associated with a sharp decline in the level of primary
production in the area.
370
Western Atlantic Bluefin Tuna (Thunnus thynnus)
The Bluefin Tuna is another species which has suffered a
catastrophic decline from over-exploitation; however,
management practices have not yet produced a recovery in
adult numbers.
The Bluefin Tuna is found on both sides of the Atlantic and
both the Eastern and Western Pacific. In the Western
Atlantic it ranges from Labrador to Brazil ICCAT, 1990).
The breeding population in the western Atlantic has been on
the decline for two decades. The population of ‘giant’, adult
fish (age 10+ years) is estimated by ICCAT (International
Commission for the Conservation of Atlantic Tunas) to have
declined by nearly 95% since 1970 (Fig. 26.6).
Because of the extremely high value of tunas (up to
US$30,000 a fish) there has been a marked reluctance to
curtail catches even when the need to do so was evident. A
total allowable catch quota was set by ICCAT in 1982 but
this was doubled in 1983 and has remained the same ever
since. Intense publicity was directed at the species in 1992
as a result of a proposal to include it in Appendix I of
CITES. The proposal was eventually withdrawn under
political pressure but a reduction in catch quotas (10%
reduction on the 1991 quota in 1992-1993 and a 25%
reduction in 1994-1995) was agreed, entering into effect in
May 1992 (ICCAT, 1991). This was expected to allow a
very slow recovery in population size but obviously not as
swift as would occur if fishing were to be halted altogether.
The amount of "illegal" catch (i.e. catch exceeding the
ICCAT quota) causes these quotas to be exceeded and
further slows recruitment.
Long-lived species, such as the Bluefin, have relative
stability in the numbers of young which survive each year
and, because of their longevity (20 years or more), they
have a steady but slow recruitment each year. In contrast,
short-lived species, such as the Atlantic Herring, have a
highly variable recruitment but are capable of recovering
rapidly. These characteristics partially explain the
differences in the success of the management programmes
for the two species but it is probable that economic factors
played a greater role. As Beverton (1991, in litt.) has
pointed out, "high prices lead to depletion, even when
abundance is low".
Future development in marine fisheries
Most fishery stocks currently used are believed to be fully-
or even over-exploited, but demand for fishery products is
predicted to rise by the end of this century (Norse, 1992).
New stocks, species, or techniques will therefore be
needed. Two further options available to increase fisheries
production are the exploitation of high seas resources, and
mariculture.
High seas resources - those over 200 miles from shore,
beyond national jurisdiction - are increasingly under
pressure from long-range fleets, which in many cases are
exploiting them as a direct result of being banned from
traditional fishing grounds in newly-formed fishery
exclusion zones. More than 400 fishery species are
Animal Use
Figure 26.4 Trends in fisheries catches, 1963-1989
100
RED
w
oO
iS
(=
fo)
~~
[=
2 60 ;—
£
= L
Lo)
=
Oo 40 —
o
= Total catch (marine and inland) Marine
ao) he
ree ee Oy Cmte en om) ae sais
=
200
[ae 1 ! 1 tI =: ! 1 | L | 1 i 1 Laer | 1 [ey | 1 St | 1 Ss
1963 1965 1967 1969 1971 1973 1975 1977 1979 1981 1983 1985 1987 1989
Year
considered to be high seas or oceanic: these include 50
species of cephalopods, 40 species of sharks, 60 species of
_ Marine mammals and 230 species of bony fish (FAO,
1990b). Most stocks of these species are dispersed and
difficult to harvest or study, and occur at much lower
densities than those in upwellings and coastal zones. The
main technique used to harvest them is pelagic drift-netting,
in which monofilament gillnets are set hanging from the
surface to a depth of 10m and left to drift overnight. This
practice causes large-scale accidental mortality of non-target
species (Norse, 1992). However, because of the concern
about the level of incidental catch, restrictions have been
progressively introduced to curb drift-netting.
The group with the greatest potential for fishery
development are the oceanic squids. Although the main
species are already fully fished or overfished, new species
and areas have development potential. In the Antarctic, krill
fisheries could be expanded, though the economic viability
of such a project is very doubtful at the moment and there
are serious problems of ecosystem management which have
yet to be solved (FAO, 1990b). Most high sea resources
under international managementsuffer from excessive effort
and depletion, and practices for responsible fishing need to
be agreed by participating nations (FAO, 1990b).
Mariculture is expected to contribute increasingly to world
fishery production. Both intensive and extensive mariculture
production have grown considerably more than capture
fisheries in the past few years. In 1988, 14.6 million
tonnes, or 14.8% of the total world catch, was obtained
from aquaculture activities, and by the year 2000 this
371
proportion may increase to 33% (Norse, 1992). Freshwater
culture of carps and other cyprinid fishes was by far the
largest component of this production, but marine species
such as mussels, oysters, salmon, shrimps and prawns were
also highly significant.
Inland fisheries
Inland fisheries (aquaculture and capture fisheries) grew
steadily by 32% over the five-year period 1984-1988 and,
in 1989, contributed 13.8% or 13.8 million tonnes of world
landings of aquatic resources (FAO, 1991a). Asia is the
major inland fisheries producer: in 1989 the continent
harvested approximately 9.5 million tonnes, 69.2% of the
world total inland catch; 98% of the Asian catch consisted
of carp and tilapia species (FAO, 1990), much of which
was produced in fish farms. Africa and the "USSR" also
produced significant quantities of freshwater fish, mainly
wild-caught, but in other continents production was
negligible in terms of volume compared to that from marine
fisheries (Fig. 26.7).
Despite their small volume relative to marine fisheries,
inland fisheries are frequently of particular subsistence
value to local communities. Aquaculture grew rapidly in the
five years 1984-1988. Global aquaculture production from
inland waters increased by 42%, with spectacular increases
in Oceania (261%) and Africa (99%). In contrast, inland
capture fisheries remained relatively stable or declined over
the same period, especially in the industrialised countries.
The general trend in the industry is thus for aquaculture to
substitute increasingly for production from capture fisheries
(FAO, 1990).
2. Uses and Values of Biodiversity
Figure 26.5 Trends in herring stocks in the past 20 years
Spawning stock blomass (1,000 tonnes)
Estimated population (1,000s)
1,500
1,000
Year
North Sea Via North Via South Irish Sea NO Spring 1S Summer
234,900
200
1970 1972 1974 1976 1978 1980 1982 1984 1986 1988 1990
Year
372
Animal Use
Figure 26.7 Catches in inland waters, 1984-1989
xe
SOON RX
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373
2. Uses and Values of Biodiversity
NON-FOOD USES
Utilitarian uses
In addition to its nutritional value wildlife may provide
important utilitarian products for both domestic and
commercial markets. Fur, hides, scales, bones, and feathers
may be used to make a variety of clothing and utensils,
while fat may be rendered for oil. Glue and household
implements such as needles and hooks can be made from
bones, scales and fins. The sale of these products on a
small or large scale can generate significant income. It is
worth noting that the exploitation of wild species does not
necessarily entail killing individual animals. For example,
the great seabird colonies of Peru are a source of guano,
used as fertilizer, while for centuries in Iceland, the down
of Eider Ducks Somateria molissima has been collected
from their nests. In Peru Vicufia Vicugna vicugna are
periodically rounded up and shorn of their extremely fine
hair, and the fibre from the Musk Ox Ovibos moschatus can
be collected from the ground during the annual moult.
Ornamental uses
Other wildlife products are valued for their ornamental,
decorative or ceremonial purposes. Elephant ivory,
tortoiseshell (derived from the Hawksbill Turtle
Eretmochelys imbricata) and furs have been much prized
and in international commerce for many centuries. Both are
now prohibited from international trade by their inclusion
in Appendix I of CITES, but ivory was formerly much in
demand for fabrication into billiard balls, piano keys and a
variety of jewellery and artefacts. Fig. 26.8 shows the
changes in the amount of ivory exported between 1979 and
1988.
The production of reptile leather has risen in importance
since the start of the present century, and demand is
continuing for manufacture of shoes and fancy goods.
Since sea turtles have been included in Appendix I, reptile
skin trade is now principally confined to three groups: the
crocodilians, lizards and snakes. Crocodilians have the
highest value skins, particularly those described by the
industry as ‘classic skins’, members of the genera
Crocodylus and Alligator. The Latin American Spectacled
Caiman Caiman crocodilus is now much more numerous in
trade, as it is in the wild, but has a cheaper skin and is
therefore well suited to the mass market.
Amongst the lizards, the tegus (genus Tupinambis) and the
monitors (genus Varanus) are large enough to produce
useful skins. Both are traded in huge volumes, up to nearly
three million skins a year for Tupinambis, almost all of
which come from Argentina. The monitor lizards can be
divided into the African species, V. niloticus and
V. exanthematicus, which are mostly imported to Europe,
the Southeast Asian species, V. salvator, which is traded to
Japan and Europe, and the Indian species, V. bengalensis
and V. flavescens, which almost all go to Japan. The last
two are included in Appendix I but are imported by Japan
under the terms of a "reservation" which enables it to
continue importing them as if they were not covered by
CITES. For this reason the statistics are incomplete.
374
The most valuable of the snakes are the large boids,
particularly the pythons, Python reticulatus, P. molurus,
and P. curtus from Southeast Asia and the African Rock
Python P. sebae. Latin American species include the
anacondas Eunectes spp. and the Boa Constrictor. Trade in
the skins of boids has remained at around three-quarters of
a million, the majority being from P. reticulatus. In order
to serve a mass fashion market, the industry has recently
been shifting away from boids towards some smaller and
cheaper species from Asia such as the Rat Snake Ptyas
mucosus and the Dog-faced Water Snake Cerberus
rhynchops. Both were formerly included in CITES
Appendix III, and were therefore incompletely recorded in
the statistics, but have recently been transferred to
Appendix II. This means that the trends shown in Table
26.11 are misleading for such species (marked with *) but
otherwise they give an overall impression of the volume and
composition of the trade. Sea snakes, especially of the
genus Lapemis and the brackish water genus Homalopsis,
are also traded in large numbers, particularly from the
Philippines, but are not included in CITES and therefore do
not appear at all in the statistics.
Most of the trade in furs derives from farms but cat (felid)
species are not farmed and the entire trade derives from the
wild. Trade in cat skins is summarised in Table 26.12.
Europe and Japan provide the main markets but North
American countries are net exporters of cat skins in most
years, the two species exported being the Lynx Felis lynx
canadensis and Bobcat F. rufa. Exports of both have
gradually declined since the early 1980s owing, principally,
to the decreasing popularity of furs as fashion items. This
shift in demand has largely been responsible for the very
marked decline in trade in cat skins from all sources.
Latin America was the main source of skins in the early
1980s, especially Paraguay, Bolivia and Argentina, but this
trade declined sharply in 1985 as a result of import
restrictions brought in by the EEC. The species in trade
were Ocelot Felis pardalis, Little Spotted Cat F. tigrina,
Margay F. wiedii and Geoffroy’s Cat F. geoffroyi. The first
three of these were transferred to CITES Appendix I in
1989 and trade in the fourth was virtually confined to old,
stockpiled skins.
Largely as a result of these legal restrictions on the supply
of the South American species, the trade has shifted in the
late 1980s to China which has been the single largest source
of skins, almost all of the one species, the Leopard Cat
F. bengalensis.
The trade in cat skins has therefore reflected major changes
in fashion, coupled with alterations in the legal control
under CITES. This has affected not only the overall volume
of trade but, possibly more importantly in biological terms,
the sources and species in trade.
Bird feathers are also used as items of adornment in many
parts of the world, often being incorporated into traditional
dress to indicate status or hierarchy. In Latin America
feathers of birds such as the Quetzal Pharomachrus
mocinno, the Roseate Spoonbill Ajaia ajaia and macaws Ara
spp. are prized for their decorative qualities. These feathers
have commanded great value throughout many generations;
Animal Use
Figure 26.8 Ivory exports from Africa, 1979-1988
OWOM Aq paiduios ‘eyep SalI :eounos
UMOYS ese suOVeU BulOdxe jueHOdUII SOW BAjOM} eUL :a]ON
BOL WO. SUOdxe AIDA! peps009d |E}0} WINWIUIW
375
2. Uses and Values of Biodiversity
Table 26.11 Approximate total net trade in reptile skins 1983—1989
TAXON 1983 1984
Chelona mydas 5,716 1,000
CROCODYLIAA spp. 0 0
Alligator mississippiensis 17,826 13,057
Caiman crocodilus 1,523,421 1,502,191
Crocodylidae spp. 0 0
Crocodylus acutus 800 0
Crocodylus cataphractus 9,911 2,030
Crocodylus johnsoni 0 157
Crocodylus niloticus 33,474 7,027
Crocodylus novaeguineae novaeguineae 30,995 30,061
Crocodylus porosus 5,495 5,839
Crocodylus siamensis 0 0
Crocodylus spp. 0 3
Melanosuchus niger 0 452
Osteolaemus tetraspis 0 20
Dracaena guianensis 51,424 71,541
Iguana iguana tt) 0
Tupinambis rufescens (0) 0
Tupinambis spp. 280,195 0
Tupinambis teguixin 1,476,779 1,858,403
Tupinambis teguixin nigropunctatus 1,254,300 799,076
Uromastyx spp. 0 0
Varanus bengalensis 0 474,491
Varanus exanthematicus 28,045 14,315
Varanus flavescens 0 56,274
Varanus niloticus 280,617 354,700
Varanus salvator 1,030,707 1,222,605
Varanus salvator cumingi (0) (+)
Varanus spp. 320 0
Atretium schistosum te) (0)
Boa constrictor ~ 143,809 32,517
Boa constrictor constrictor 17,004 15,111
Boa constrictor occidentalis 0 0
Boidae spp. 0 0
Cerberus rhynchops * 0 19,250
Eryx muelleri (0) 0
Eunectes murinus 9,842 25,331
Eunectes notaeus 17,883 44,376
Morelia spilota (0) 0
Morelia spilota spilota 0 0
Naja naja * te) 2,463
Ophiophagus hannah * 0) 0
Ptyas mucosus * (0) 712,671
Python curtus 43,929 42,204
Python molurus 155 0
Python molurus bivittatus 117,475 156,486
Python regius 0 0
Python reticulatus 478,901 591,168
Python sebae 1,047 782
Python spp. 215 945
Sanzinia madagascariensis () (0)
Vipera russelii 0 0
1985 1986 1987 1988 1989 AVERAGE
tt) tt) 0 0 t) 5,575
0 t) 0 0 2 2
13,228 24,023 33,080 38,720 63,860 32,290
1,738,423 862,059 639,780 882,305 238,409 1,155,371
17 t) () ts) 0 17
t) t) t) 0 59 173
0 11 149 1,193 570 2,348
0 t) 826 t) 614 167
9,378 18,753 24,156 29,524 35,415 23,376
56,924 43,972 39,340 38,171 23,768 35,430
9,160 7,340 8,431 10,259 6,008 7,400
351 605 2,132 2,050 t) 1,219
21 0 5 t) t) 4
(0) te) C0) tt) 0 452
38 41 i) 28 224 63
0 26,639 i) 0 t) 35,667
t') te) 18,755 179 t) 9,204
4,300 0 147,687 1,047,429 1,196,104 223,778
t) 194,190 96,913 96,657 86,956 119,842
1,480,744 1,393,044 1,466,912 873,625 817,675 1,440,234
166,842 33,894 7,001 0 0 990,163
t) 40,000 t¢) t) 0 40,000
296,684 24,040 0 0 0 76,687
144,460 44,230 4,297 76,461 7,212 40,941
196,316 t°) 0 te) 0 69,055
444,295 302,747 712,997 722,532 651,753 507,848
1,218,145 1,215,784 1,880,726 1,614,836 1,540,980 1,436,327
t) 15,996 6,951 29,167 0 13,973
0 1,734 16,991 1,374 ts) 9,310
te) 0 0 28,290 tt) 28,290
20,889 25,591 4,919 1,443 2,125 37,785
te) 0 688 te) te) 16,211
0 t) 901 Oo t) 901
tt) 58 530 t) te) 59
204,224 821,964 700,072 771,428 38,604 295,753
0 1 0 t°) t) 1
10,643 6,829 10,759 15,505 2,081 12,021
22,483 19,110 7,160 19,839 0 21,348
0 0 () 13 0 13
0 0 () 3,000 t) 3,000
8,966 266,600 62,510 77,009 30,593 24,440
662 1,455 320 t°) t) 651
819,055 2,658,195 1,957,577 1,615,906 1,061,874 1,531,584
0 83,922 77,293 172,203 29,923 71,244
te) t) t) t) t¢) 155
211,414 45,843 70,844 55,132 20,054 112,559
17 136 t°) 152 88 78
539,265 569,084 736,847 767,272 456,419 593,684
2,494 19,735 15,569 64,594 9,666 16,686
38 3,508 57 t) 1,205 1,587
1 t) (0) t) 0 1
te) 227,163 22,303 42,189 17,485 112,551
Note: * indicates species which were moved from Appendix III of CITES to Appendix II within the period 1983 —89 and have thus been incompletely
recorded in statistics.
Source: Annual reports of Parties to CITES, compiled by WCMC.
indeed the Incas collected tribute in exotic bird feathers
from their Amazonian subjects. The trade in feathers to
supply Western fashions, which was at its height in the
early 1900s, has largely disappeared, but was once of great
commercial significance. Between 1899 and 1920 over
15,000kg of egret and heron feathers, representing plumes
from an estimated 15-20 million birds, were exported from
South America (Redford and Robinson, 1991). The Ostrich
farming industry, established in South Africa but having
now spread to the USA and other African countries, was
originally almost exclusively for the feather trade, but skins
and meat are now more economically important.
The butterfly trade is largely based on ornamental use with
some research and education value. Butterflies of the family
Papilionidae feature prominently in trade. This family
376
contains the spectacular ‘birdwings’ which include the
world’s largest species of butterfly, Queen Alexandra’s
Birdwing Ornithoptera alexandrae with a wingspan of up to
250mm. The large butterflies are generally traded as
individual items and some are ranched in order to produce
perfect intact specimens. Many of the most spectacular and
endangered species have various levels of protection under
CITES as well as under local legislation. There is also a
major trade in less spectacular tropical species for
incorporation in ornaments and souvenirs. This latter trade
centres on Taiwan, where estimates suggest between 15
million and 500 million butterflies are traded annually
(Pyle, 1981). There has been little monitoring to assess
whether this trade is sustainable. Another more recent trade
involves live butterflies which are transported (most often
now as pupae) to provide exhibits at a variety of locations,
Animal Use
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377
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Table 26.12 Net imports and exports of cat skins, 1980—1989 (continued)
NET EXPORTS
NET IMPORTS
1981 1982 1983 1984 1985 1986 1987 1988 1989
1043. 2127 «#61179 «#491887 §=1789
1980
1989
1981 1982 1983 1984 1985 1986 1987 1988
218 158
1980
9035 772 962
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75
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379
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116
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Somalia
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OTHER
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119
104-1008
164
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105
1309
410
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Animal Use
104 1008 119
164
692
57
105
1309
Source: Annual reports of Parties to CITES, compiled by WCMC
Country Unknown
2. Uses and Values of Biodiversity
such as zoos and butterfly houses in various parts of the
world. The livestock for display is mainly captive bred at
butterfly farms in various tropical locations.
Among invertebrate-based products, the silk industry
produces a luxury commodity on a massive scale and is of
Major importance to many countries with rural-based
economies. Most world trade uses the product of one
species, the Mulberry Silk Moth Bombyx mon
(Bombycidae), a domesticated species. The silk produced by
this species is the finest quality of those available. Other
species which contribute to the world trade are mostly from
the family Saturniidae. These are all known as ‘wild’
silkworms, even though the eri (Attacus ricini), tussah
(Antheraea spp.) and muga (A. assama) silkworms are
partially domesticated. Sericulture (silkworm husbandry) is
an important source of revenue for many people. Table
26.13 shows the main countries producing raw mulberry
silk and silk waste and their estimated 1989 production. The
process of silk production is very efficient in that little is
wasted. The high-protein byproducts of sericulture
(mulberry leaves and silkworm rearing litter) make good
animal feeds; the moth pupae killed after spinning their
cocoons can be used as human or animal food and are also
used in soap and cosmetics; and the mulberry trees which
are cultivated to feed the silk worm can be used for timber
and will often grow on areas of land unsuitable for other
agriculture (Greenhalgh, 1986).
Other important insect products include lac, from Laccifer
lacca and cochineal, from Dactylopius coccus, both of
which are Homopteran scale insects. Lac is refined to
produce shellac, widely used as a base for polishes and
other wax products, with an estimated US$ 9 million used
in the USA in 1981 (Lindberg, 1988). Cochineal, a brilliant
red colouring agent, formerly widely used in the food and
cloth industries, is commercially obtained from a semi-
domesticated form of the insect. The cochineal trade was
very much reduced by the invention of synthetic dyes and
today only Peru and the Canary Islands still produce the dye
for export.
Medicinal and biomedical uses
Animal products are widely used in medicines by traditional
societies, and even urbanised societies may retain their faith
in traditional animal-based remedies, particularly in Asia.
For example, Levy-Luxereau (1972) described 181 animal-
based remedies used by the Hausa tribe in Niger, most of
which were derived from wild species. In local markets in
Brazil, dried lizards of several species, the genitalia of
dolphins, fox fur, and many other pieces of wild animals
are sold for medicinal and magical purposes (Redford and
Robinson, 1991). Medicinal wildlife products are frequently
traded internationally particularly to satisfy the demand for
traditional oriental medicine, a trade which can be
extremely lucrative. International trade in the medicinal
products of deer, especially antlers, tendons and musk was
calculated to be worth some US$30 million a year, almost
as much as the international trade in meat (Luxmoore,
1989). Some products are so valuable that the trade can
continue even when the species become extremely rare in
the wild and, in such cases, it may pose a severe threat to
their survival. Trade in musk (from musk deer Moschus
380
spp.), bears’ gall bladders, tiger bones and, most notably,
thino horn have all been blamed for the decline in
populations of several species.
The use of leeches for medical purposes probably began in
India but was first described in writing by the Greeks in the
2nd century BC (Conniff, 1987). Leeching became very
fashionable in Europe during the mid-19th century to the
point where the European Medicinal Leech Hirudo
medicinalis is now threatened in the wild and included in
CITES Appendix II. Present day usage of the live animal is
now mostly restricted to micro-surgery where the sucking
action and substances produced by leeches during feeding
help survival of accidentally severed parts, such as fingers
and ears, after re-attachment. Leech saliva contains
anticoagulants, anaesthetics, vasodilatory agents and a
spreading factor (which allows the other agents to spread
far beyond the edges of the incision) all of which have
potential uses in a range of research and medical fields. For
instance, Hirudo medicinalis is used as a source for hirudin,
an anticoagulant which can help prevent blood clots from
forming. The Giant Amazon Leech Haementeria ghilianii
uses a different chemical, hementin, for a similar purpose.
However this factor is not only capable of preventing
clotting, it can dissolve already formed clots. The spreading
agent is also produced commercially as are several other
leech-derived products. Leeches of various species are
farmed commercially in the UK to supply these different
outlets.
Horseshoe Crabs (Limulus polyphemus) are extremely
ancient in evolutionary terms and possess several unique
features. They have been used in fundamental research into
vision and the clotting property of Horseshoe Crab blood is
exploited in human blood testing. Gram-negative bacteria
are responsible for a wide range of serious diseases in man,
such as spinal meningitis and gonorrhoea, and are
sometimes responsible for contamination of manufactured
drugs. The blood of Horseshoe Crabs clots rapidly as soon
as it comes into contact with gram-negative bacteria or their
endotoxins. Refined and freeze-dried samples of Horseshoe
Crab blood can therefore be used in a very accurate assay
for the presence of these endotoxins, allowing rapid
diagnosis of disease and routine checking of purity of drug
samples. These tests have largely replaced less accurate
assays carried out on rabbits. Although blood is obtained
without killing the animals, which are collected, bled and
released, there has been some concern over the long-term
impact of this practice on Horseshoe Crab populations.
Substances with great potential uses in medicine have also
been isolated from snake venoms; these include coagulating
enzymes, anticoagulants, neurotoxins and cytotoxins. The
first two have been used in the study and control of
bleeding disorders in man and the development of fast and
accurate assay methods to test prothrombin in human blood,
so helping prevent thrombosis, or blood clot formation.
Neurotoxins may be useful as anaesthetics, and cytotoxins
could prove useful in cancer treatments.
An important use of live animals, particularly primates, is
as experimental animals in the biomedical trade. The total
volume of trade in 1989 was about 42,000 (Table 26.14),
the majority again being imported to Europe and the USA.
Table 26.13 Silk production:
ASIA
Afghanistan 60
Cambodia 14
China 42,044
India 10,500
Iran, Islamic Rep 850.
Japan . 7,000
Korea, Dem People’s Rep 3,000
Korea, Rep 1,400
Lebanon 6
Thailand 1,250
Turkey 250
Viet Nam 450
USSR 4,400
Animal Use
FAO estimates for 1989 in tonnes
EUROPE
Bulgaria 160
Greece 11
Italy 20
Poland 3
Romania 185
Spain 15
Yugoslavia 45
SOUTH AMERICA
Brazil 1,900
AFRICA
Egypt 11
Madagascar 15
Source: FAO, 1990a,b. FAO Yearbook: Production 1989. Food and Agriculture Organisation of the United Nations, Rome.
Note: All figures are estimates.
The main exporting countries are Indonesia and the
Philippines, most of the exports comprising the single
species, the Cynomologus Macaque Macaca fascicularis.
The same species occurs in large feral populations in
Mauritius, and this small island constituted the world’s third
largest exporter of live primates. In the past, South America
has been a major source of primates but this has now
declined owing to trade restrictions and any residual need
is filled by captive breeding. The species most commonly
exported were Common Marmosets Callithrix jacchus and
Squirrel Monkeys Saimiri sciureus, followed by marmosets
of the genus Saguinus, and Night Monkeys Aotus trivirgatus
(Redford and Robinson, 1991).
Chimpanzees Pan troglodytes are also widely used in
biomedical research, and in 1990 approximately 1,300 were
held by biomedical facilities in USA alone. International
trade is now banned by the inclusion of the species in
CITES Appendix II but illegal capture of Chimpanzees from
the wild and export from Africa continue, driven by the
high market value of infants - around US$25,000. This
trade is blamed for the continuing decline of wild
populations (CCCC, 1990).
Increasingly, primates are being bred in captivity for
biomedical research. This is preferable as it does not risk
depleting wild populations and it produces a genetically
uniform and disease-free stock. Some species are only bred
in very small numbers on an experimental scale, but others,
such as M. fascicularis and C. jacchus, are bred
commercially.
Working animals
A number of wild animal species are trained to assist in
various human activities. For example, the Indian Elephant
Elephas maximus has been used for centuries as a draught
animal in forest industries, warfare, and for ceremonial
purposes. There are currently approximately 16,000 tame
elephants, the majority employed in the timber industry in
India, Myanmar and Thailand. Although the species breeds
well in captivity most working elephants are captured from
the wild and subsequently tamed. Both otters Lutra spp. and
381
cormorants Phalacrocorax spp. are used for fishing in
China and Southeast Asia. After training, the animals are
fitted with restraints to prevent them damaging or
swallowing the fish they catch (leather straps over the
canines in the case of otters, neck rings on the cormorants),
and generally kept tethered to the fisherman’s boat during
fishing. Pig-tailed Macaques Macaca nemestrina are often
captured in Southeast Asia and trained to climb trees and
throw down ripe fruit and coconuts.
Trained wild animals are also widely used in sport hunting.
Falconry, the sport of using falcons, hawks and sometimes
eagles to capture and kill wild game, relies on the training
of both young and old birds taken from the wild. Practised
worldwide, it is still a popular pastime in India, Pakistan
and Saudi Arabia. The Cheetah Acinonyx jubatus was once
used by wealthy Indians as a trained courser, and might
well have become domesticated if it had not been for its
reluctance to breed in captivity.
Mutually profitable associations can even arise between men
and totally untrained wild animals. Thus some Amazon
Indians have developed close fishing partnerships with
individual Amazon River Dolphins Inia geoffrensis. The
man calls his dolphin by whistling and it feeds on the
opposite side of the river from his canoe. The dolphin’s
activities drive the fish towards the man, and vice versa,
resulting in more successful fishing for both. The
association between honey guides Indicator spp. and men,
in which the bird leads the hunter to wild bees’ nests by
calling insistently and fluttering its wings in return for a
share of the spoils of honey, wax and grubs, is another
example (Barton, 1986).
Pollination
A vital operation carried out by insects, mostly
independently of man, is pollination. Many of man’s
most important crops rely on pollination by insects, and of
these insects bees are by far the most important. The bees
fall into two categories, wild bees and domesticated
honeybees. Honeybees of the genus Apis have been
cultivated by man for many centuries in various regions of
2. Uses and Values of Biodiversity
Table 26.14
Live reptile and primate trade in 1989
LIVE REPTILES LIVE PRIMATES
IMPORTS! EXPORTS' IMPORTS‘ Exports!
WORLD 438875 386754 40619 42249
ASIA 32011 57297 4815 26018
Bangladesh is) te) 8 i)
Brunei i) 0 4 0
China 17 te) t) 1292
Cyprus 43 te) te) )
Hong Kong 135 i!) i) 24
India 2 is) 29 0
Indonesia te) 7914 i) 16501
Iran te) te) 180 t)
Israel 30 69 0
Japan 30623 () 4184 ()
Jordan te) 6 0 0
Korea, Rep 2 i) 2 0
Laos te) 150 it) 44
Lebanon i) 0 1 0
Malaysia 0 19708 13 0
Myanmar 0 0 0 56
Pakistan 0 1 0 0
Philippines te) 19059 () 8963
Saudi Arabia 10 te) 53 t)
Singapore 1965 () () 32
Sri Lanka 2 0 0 2
Taiwan t) 4 223 0
Thailand te) 2413 28 0
Turkey i) 7504 0 0
United Arab Emirates 112 0 21 i}
Viet Nam i) 501 ts) 4
Yemen 0 7 i?) 0
USSR (former)
ty) 52121 1630 0
EUROPE 131561 224 14626 3
Austria 32252 0 135 0
Belgium 1270 0 1143 0
Czechoslovakia 0 146 35 0
Denmark 29 ie) 60 1)
Finland 2 (e) () 1
France 31522 0 2333 t)
German Dem Rep 225 te) 13 0
Germany, Fed Rep 23745 () 190 tt)
Greece te) 29 3 0
Hungary 591 () 85 ()
Ireland 2 0 t) 1
Ital 3692 te) 1169 ()
Mal 10 te) ts) tt)
Monaco 254 te) 3 tt)
Netherlands 16912 0 2264 ()
Norway 5 0 0 1
Poland i) 40 1 tt)
Portugal te) te) 60 t)
Romania 9 i) 18 0
Spain 1706 0 76 0
Sweden 97 ts) 698 0
Switzerland 6912 is) 111 t)
United Kingdom 12326 te) 4183 0
Yugoslavia tt) 9 2046 0
NORTH & CENTRAL AMERICA 272038 128416 20014 1515
Antigua 0 180 () tt)
Bahamas 2 te) tt) LY)
Barbados () 0 ts) 986
Belize 0 4 () 0
Canada 5286 () 1350 (')
Cayman Islands tt) 19 0 0
Costa Rica 0 48 t) tt)
Dominican Rep 0 11 20 0
El Salvador 0 8801 te) 0
Haiti 0 5635 () ()
Honduras ts) 112625 ts) 528
Mexico 0 12 221 0
Netherlands Antilles 0 75 0 0
Nicaragua te) 1000 te) 0
Panama ts) 2 te) 0
Trinidad and Tobago te) 4 ts) 1
United States 266750 te) 19323 0
SOUTH AMERICA 120 57427 110 3501
Argentina 0 2183 0 91
Bolivia tt) 0 i) 5
Brazil 105 i) 0 231
Chile tt) 903 47 0
Colombia () 7985 () 1
Ecuador 15 i) tt) 2
French Guiana tt) ts) 63 0
Guyana 0 9842 0 2822
Peru 0 9765 t) 342
Suriname t) 26749 ) 3
Venezuela 0 te) 0 4
382
Table 26.14
Animal Use
Live reptile and primate trade in 1989 (continued)
LIVE REPTILES LIVE PRIMATES
IMPORTS! EXPORTS! IMPORTS! EXPORTS!
OCEANIA 272 3142 2 272
Australia 272 0 0 272
Fiji (0) (:) 2 0
New Zealand to) 1 0 ts)
Papua New Guinea 0 1 0 ts)
Vanuatu (0) 16 0 0
Solomon Islands 0 3124 0 ts)
AFRICA 1813 140157 7 9613
Angola tt) tt) 3 0
Benin i) te) 0 1
Botswana it) 671 1 0
Burkina Faso te) te) 2 0
Burundi te) 4326 (¢) te)
Cameroon 0 te) te) 80
Comoros 0 428 () 0
Congo 0 2 1°) 4
Céte d'ivoire ts) 12 tt) 0
Egypt () 1060 () ()
Ethiopia (0) 0 0 492
Gabon 0 () 0 2
Ghana tt) 25400 te) 249
Guinea 0 6 ts) 2
Kenya tt) 53 te) 2176
Lesotho to) 1462 0 ts)
Liberia 0 150 0 ()
Libya 12 (0) 2 0
Madagascar 0 24901 (0) 4
li (e) 10 ts) ()
Mauritius 388 0 ts) 3215
Morocco t) 15 4 tt)
Mozambique 0 368 ) 0
Namibia 0 1028 1 t)
Niger (e) (e) te) 2
Nigeria 0 2 0 0
Reunion 19 (0) ts) ()
Senegal te) te) 0 607
Seychelles 0 176 0 0
South Africa 1368 0 0 13
Sudan 0 2 (0) {0}
Swaziland 0 0 2 0
Tanzania 0 4416 i) 2387
Togo 0 74975 (0) 363
Tunisia 26 () 0 3
Uganda t) 0 1 0
Western Sahara 0 126 0 0
Zaire 0 226 te) 12
Zambia 0 195 1 te)
Zimbabwe 0 147 0 1
OTHER 160 91 135 427
Notes: ' figures are net.
Sources: Annual reports of Parties to CITES compiled by WCMC.
the Old World where they occur naturally. In the Americas
where there are no native Apis, ‘stingless’ bees of the
family Meliponidae have traditionally been kept, but the
more recently introduced A. mellifera from Europe has
become the main species in the beekeeping industry. A
main reason for the domestication of bees has always been
the production of honey and beeswax (Table 26.5).
However, the importance of these insects for pollination has
not been missed and apiculture has usually gone hand-in-
hand with agricultural production of crops requiring bee
pollination (see Table 26.15).
Wild bees of hundreds of species are also important
pollinators and are more effective than the honeybees for
certain crops and in colder climates.
Sport hunting
In many societies animals are hunted for pleasure and, in
affluent societies, private individuals may pay large sums of
money for the privilege. In many cases the offtakes are
controlled by a system of hunting licences or permits sold
by the government, which can raise significant revenues for
central or local treasuries. For example, deer, gamebird and
383
wildfowl harvests are regulated in this manner in North
America. Owners of private land, including in some African
countries, are able to sell the right to shoot animals on their
land to visitors, considerably enhancing the economic value
of wildlife.
Recreation, tourism, aesthetic value
Wildlife also has enormous recreational and aesthetic value.
Many people derive pleasure from wildlife either by
observing them in the course of their daily lives, by making
special excursions to view them, by watching them on film
or television, or simply by knowing that they continue to
exist, without necessarily wishing to see for themselves.
This kind of non-consumptive use is very difficult to
evaluate but it is possibly the single greatest economic value
of wildlife. One of the easier techniques is to quantify what
tourists are prepared to pay to observe wild animals in their
natural habitat. This is of considerable interest in
developing countries because overseas tourists bring in
much-needed foreign exchange. It has been estimated that
visitors pay almost US$200 per person to spend an hour
with wild but habituated Mountain Gorilla Gorilla gorilla
beringei groups in Rwanda. These visits generate nearly
2. Uses and Values of Biodiversity
Table 26.15 Selected crops of commercial importance in the EC for which there is
agreement of the importance of insect/bee pollination from several sources
FAMILY GENUS AND SPECIES COMMON EC PRODUCTION REPORTED NEED FOR INSECT
NAME (x1000 tons) VISITORS POLLINATION
D P
Compositae Helianthus annuus Sunflower 3,908 H,B,S 1.0 0.9
Cruciferae Brassica campestris Turnip rape ? H,B,S, - -
Cucurbitaceae Citrullus lanatus Water melon 1,838 H,S 0.7 0.9
Cucumis sativus Cucumber/gherkin1,372 H,S 0.9 0.9
Cucumis melo Melon 1,654 H 0.8 0.9
Lauraceae Persea americana Avocado 29 H,S 1.0 0.9
Leguminosae Medicago sativa Lucerne/alfalfa ? H,B,S 1.0 0.6
Phaseolus multiflorus Runner bean ? H,B - -
Trifolium pratense Red Clover ? H,B,S - -
Rosaceae Prunus amygdalus Almond 347 H > 1.0 1.0
Prunus sp. Cherry 546 H,B 0.9 0.9
Prunus sp. Pear 2,631 H,B 0.5 0.9
Pyrgus malus Apple 9,321 H,B,S 1.0 0.9
Rubiaceae Coffea sp. Coffee ? H,S - -
Source: Compiled from several tables in : Corbet, S.A., Williams, I.H., and Osborne, J.L. 1991. Bees and the Pollination of Crops and Wild
Flowers: changes in the European Community. Review commissioned by Scientific and Technical Options Assessment, European Parliament. These
tables are themselves drawn from a number of studies, some carried out in the USA.
Notes: ? Statistics unknown. H = honeybee. B = bumblebee. S = solitary bee. D = proportion of yield attributable to insect pollination. P =
proportion of effective insect pollinators that are bees.
US$1 million per year in direct park revenues (Vedder and
Weber, 1990). In Kenya, the African Elephant was
estimated to be worth some US$25 million a year to the
tourist industry (Brown, 1989). The enjoyment of wildlife
and wild habitats is one of the tourist industry’s most
rapidly expanding sectors. Thus it is primarily the wildlife
which draws people to places such as East and southern
Africa, the Galapagos, the Valdés Peninsula and the
Brazilian Pantanal.
Social and cultural significance
Wildlife influences the philosophy, language, art, religion,
and social structure itself of many societies. In African
cultures wild animals figure prominently in animist beliefs,
mythology, and works of art such as carvings and paintings.
Economic dependence on wild resources produces a close
relationship between the ecological factors governing
wildlife and social organisation in some tribes. For
example, seasonal alterations in hunting technique
(individual hunting in the wet season, cooperative netting of
game in the dry season) in the Babinga people of the
Central African Republic affect the location of
encampments, seasonally influencing group composition,
social rapport and material exchange. In many cultures a
man’s social worth is measured by his prowess as a hunter.
Companion animals
Wild animals have been captured and kept in captivity for
a variety of reasons for centuries, including as pets or
companion animals, for entertainment, and for private and
public display in zoos and menageries. In South America,
a survey of four Kayapo villages revealed that at least 31
species of animal were kept as pets, including five species
of turtle, 16 species of parrot and macaw, a lizard and a
spider (Redford and Robinson, 1991). Pet-keeping is an
almost ubiquitous human activity, and there is a thriving
export trade in wild animals for pets from many tropical
countries to the developed countries, involving large
384
numbers of species and significant sums of money. For
example, in the USA the annual retail turnover of parrots
alone, both wild and captive-bred, has been estimated at
US$300 million (Hemley, 1988). The gross retail value of
parrots exported from Neotropical countries from 1982 to
1986 was estimated to be some US$1.6 billion. Parrots are
more valuable than the smaller species of birds used in the
pet trade and, although they are traded in smaller numbers,
they contibute a disproportionate amount to the total value
of the trade. Bird imports to the USA from Indonesia in
1986 and 1987 were worth US$4.4 million, an average of
US$79 a bird, while the larger numbers of birds imported
from Senegal over the same period were only worth an
average of US$1.37 each because they comprised mostly
the smaller seed-eating species (Thomsen et al., 1992).
Table 26.16 shows a summary of the trade in live parrots
from 1980 to 1989 recorded in CITES annual reports. The
overall levels of trade appear to have increased from 76,629
in 1980 to a peak of 625,799 in 1988 before declining
slightly in 1989. This finding must be treated with some
caution because the standard of reporting of trade has not
remained constant over this period. The number of Parties
to CITES has increased from 1980 to 1989 and this will
almost inevitably entail an increase in the volume of trade
reported. Furthermore, the ability of Parties to monitor
their trade has improved as more sophisticated mechanisms
and procedures have been set up. Thus, although France
has been a Party to CITES since 1978, its reported volume
of parrot imports has climbed from zero in 1980 to 34,643
in 1989. There may have been some increase in the volume
of trade over this period but the major increase, apparent
between 1984 and 1985, was the result of the adoption of
new procedures to report on imports of Appendix II
species. Nevertheless, data from a range of countries
suggest that there has been a genuine increase in trade over
the period 1982-1988. The apparent decline in the volume
of trade in 1989 is probably attributable to the delay in
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submitting annual reports. Table 26.16 shows that Europe
and North America are the largest importers of live parrots
and that the main exporters are in South America, Africa
and Asia. The principal individual countries are Argentina,
Indonesia, Tanzania and Senegal.
Reptiles are also used in the pet trade, the total volume of
specimens recorded in CITES annual reports approaching
half a million specimens in 1989 (Table 26.14). Over half
of the total was imported by the USA and the majority of
the remainder by countries in Europe. The main source
countries were in Africa and Latin America.
DOMESTIC LIVESTOCK
The development of settled agriculture and animal
husbandry has enabled human societies to live at high
population densities which is a prerequisite for cultural
development of the kind based on extensive division of
labour. Consequently, hunter-gatherers everywhere have
been displaced by agriculturists and there has been an
adaptive radiation of domestic mammalian livestock into a
greater range of habitat types than those occupied by any
wild mammal. When ecological conditions have worsened,
farming systems have been adapted, usually by adopting
seasonal patterns of exploitation (pastoralism).
Domestication and breeds
In the process of domestication small numbers of wild
animals were enfolded into human societies, which assumed
responsibility for them and exerted control over their
breeding. Almost all major domestication took place in
western Asia and the Near East, from about 10,000 BC.
A domestic animal can be defined as one "that has been
bred in captivity for purposes of economic profit to a
human community that maintains complete mastery over its
breeding, organisation of territory, and food supply"
(Clutton-Brock, 1987). These truly domestic (or ‘man-
made’) animals may differ radically from their wild
ancestors in respect of a variety of features. Animals
generally regarded as domestic are listed in Table 26.17.
A second group includes domesticated (or ‘exploited
Captive’) animals, which are tamed or conditioned
individuals from populations whose breeding is not wholly
under human control and which, as a result of continuing
natural selection, tend to retain features of value in
demanding environments. Principal domesticated vertebrates
are listed in Table 26.18.
These categories are somewhat arbitrary in so far as
domestic animals (in the broad sense) represent a wide
spectrum of conditions and particular cases, but the
distinction can be useful. There is a clear difference
between traditional European farm livestock on the one
hand, and reindeer, yak and dromedary on the other. In the
latter group, whilst there may be some artificial selection
for particular traits (for pack or racing dromedaries, for
example), artificial selection has been limited because the
animals’ continuing close adaption to particularly harsh
environments is the feature of special value to humans.
389
Animal Use
However, several intermediate groups could, with emphasis
on different aspects of breeding or husbandry, be regarded
as either truly domestic or ‘exploited captive’. Semi-
domesticated animals are thus more difficult to define as
they range from those such as the Silver Fox, which has
been bred for many generations to produce distinctive pelt
colours, to species, such as the musk deer, which are only
kept on a very small, and at present experimental, scale.
Pets make up a further group of animals that is extremely
important in many if not most societies and pet keeping
could have been a first stage in domestication (Serpell,
1989).
It is important to distinguish animals of the above kinds
from tamed animals; as Clutton-Brock (1987) points out,
any young mammal, taken from its mother, can be tamed
but this is not necessarily a permanent state. Whether or not
the tameness persists into adult life depends on the species,
and any offspring would have to be tamed in their turn.
Domestic species of ungulate (hoofed mammals) tend to be
large (over SOkg) non-selective feeders whose native habitat
is open terrain or mountains (Tennessen and Hudson,
1981). The smaller-bodied species from habitats like forests
and swamps (including most deer and antelopes) include
several semi-domesticated species but no fully domestic
ones, apart from the pig. Generally, domestic ungulates are
non-territorial, living in groups of 15-100. It is not clear
why other ungulate species, such as the Eland Taurotragus
oryx and the European Bison Bison bison were never
domesticated. Perhaps human societies sharing their ranges
obtained enough food by hunting and there was not the
population pressure that seems to have been the impetus for
domestication of other ungulates.
The changes in mammals consequent on domestication are
physical and behavioural, and have a genetic basis
(Clutton-Brock, 1987). Body and brain sizes were reduced
and proportions altered. Ears were lengthened in most
species (except the horse), and the tail was lengthened in
sheep, and selected to be curly in pigs and dogs. There was
greater variability in the pelage. Particularly in the dog and
pig, but also in the Niata cattle of Uruguay and to some
extent in Jersey cattle (Darwin, 1868), the facial region and
jaws were shortened. The teeth were reduced in size and
became more crowded, especially in the dog. Greater
docility, and changes in vocalisations, particularly in the
dog, were accompanied by a retention of juvenile patterns
of behaviour, particularly playfulness. Breeding seasons
were lengthened, but less so in the most primitive breeds,
e.g. of dogs and of sheep (Brisbin, 1977; Lincoln, 1989).
While humans selected for behavioural traits and visible
markers like coat colours, the animals were also subject to
local pressures of natural selection.
It was the interaction of natural (environmental) selection
and human (artificial) selection that led to distinct breeds.
A breed can be defined as, "a group of animals that has
been selected by man to possess a uniform appearance that
is inheritable and distinguishes it from other groups of
animals within the same species" (Clutton-Brock, 1987).
Wild animals are adapted to the physical and biological
environments, and domestic animals are also subject to
2. Uses and Values of Biodiversity
Table 26.17 Domestic livestock
DOMESTIC FORM
MAMMALS
LAGOMORPHA
Rabbit Oryctolagus
cuniculus
RODENTIA
Guinea pig Cavia porcellus
CARNIVORA
Dog Canis familiaris
Ferret Mustela furo
Cat Felis catus
PERISSODACTYLA
Horse Equus caballus
Donkey Equus asinus
ARTIODACTYLA
Pig Sus domesticus
Llama Lame glama
Alpaca Lama pacos
Dromedary Camelus
dromedarius
Camelus bactrianus
Rangifer tarandus
Bactrian camel
Reindeer
Water buffalo Bubalus bubalis
Cattle (taurine) Bos taurus
Cattle (zebu) Bos indicus
Yak Bos grunniens
Mithan Bos frontalis
Bali cattle Bos javanicus
Goat Capra hircus
Sheep Ovis aries
BIRDS
GALLIFORMES
Chicken Gallus gallus
Turkey Meleagris gallopavo
ANSERIFORMES
Goose Anser anser
Chinese goose Anser cygnoides
Muscovy duck Cairina moschata
Mallard duck Anas platyrhynchos
COLUMBIFORMES
Pigeon Columba livia
INSECTS
Honey bee Apis mellifera and
other Apis spp.
Silk worm Bombyx mori
Silk worms - e.g. Antheraea
other semi- pernyi, A. mylitta,
domesticated Attacus ricini,
species Anaphe spp.
Cochineal bug Dactylopius coccus
WILD PROGENITOR
European rabbit 0. cuniculus
Cavia aperea
Wolf Canis lupus
Polecat Mustela putorius
Steppe polecat Mustela eversmanni
Wild cat Felis silvestris
Wild horse Equus ferus
African ass Equus africanus
Wild boar Sus scrofa
Guanaco possibly Lama
guanicoe
Lama sp.
Dromedary Camelus sp.
Camelus ferus
Rangifer tarandus
Bactrian camel
Reindeer
Water buffalo Bubalus arnee
Aurochs Bos primigenius
derived from 8. taurus
Yak Bos mutus
Gaur Bos gaurus
Banteng Bos javanicus
Wild goat Capra aegagrus
Mouflon Ovis orientalis
Red junglefowl Gallus gallus
Ceylon Gallus lafayetii
junglefowl
Grey junglefowl Ga//us sonneratii
Green Gallus varius
junglefowl
Wild turkey Meleagris gallopavo
Greylag goose Anser anser
Swan goose Anser cygnoides
possibly B.
mandarina
Dactylopius coccus
FIRST
KNOWN DOMESTICATION
DATE PLACE
36 BC S Europe
1000 BC S America
12,000 BC Iraq
20 AD S Europe
1600 BC Egypt
3500 BC S Ukraine
4000 BC Egypt
7000 BC W Asia
5500-4200 Andean
BC plateau
3000 BC W Asia
3000 BC W Asia
? ?
not known China/Indo-
China
6200 BC Turkey
not known not known
2500 BC
7-8000 BC W Asia
7-8000 BC W Asia
4000 BC S and SE
Asia
1500 AD Europe
500 BC
500 BC
500 BC
3000 BC
2000 BC
2500 BC Asia
pre 1200AD Mexico,
Central
America
DISTRIBUTION OF WILD
PROGENITOR
SW Europe, possibly N Africa
S America
N hemisphere
Europe
USSR, China
Europe, Asis, Africa
Russia, Central Asia
N Africa, possibly W Asia
Europe, Asia and N Africa
S America
S America
Asia, possibly N Africa
Russia, Central Asia
Arctic, sub-Arctic (feral:
Greenland, Iceland, S
Georgia)
India, S Asia, possibly W
Asia
Europe, Asia, N Africa
Tibet, Himalayas
S and SE Asia
SE Asia including Borneo
W Asia
W Asia
S and SE Asia
C -N America
N Europe, N Asia to NW
Africa
Europe, Asia, N America, N
Africa
Mexico to Peru and Uruguay
Europe, Asia, N America, N
Africa
Europe, N Africa, India to
Japan
Africa, Europe
Americas
Source: compiled from various sources; mammal data after Clutton-Brock, J. 1987. A Natural History of Domesticated Mammals.
390
Table 26.18
NAME LATIN NAME
DEER
Elk/Moose Alces alces
Axis Deer Cervus axis
Fallow Deer Cervus dama
Red Deer/Wapiti Cervus elaphus
Sika Deer Cervus nippon
Rusa Deer Cervus timorensis
Sambar Cervus unicolor
Pére David's Deer
Dwarf Musk Deer
Elaphurus davidianus
Moschus berezowskii
Himalayan Musk Moschus chrysogaster
Deer
OTHER UNGULATES
Impala Aepyceros melampus
Springbok Antidorcas marsupialis
Blesbok Damaliscus dorcas
Grant’s Gazelle Gazella granti
Thompson’s Gazella thomsoni
Gazelle
Fringe-eared Oryx Oryx beisa
Gemsbok Oryx gazella
African Buffalo Syncerus caffer
Eland Taurotragus oryx
FUR-BEARERS
Arctic Fox (Blue Alopex lagopus
phase)
Long-tailed Chinchilla laniger
Chinchilla
Sable Martes zibellina
Polecat/Fitch Mustela putorius
Mink Mustela vison
Coypu Myocastor coypus
Racoon Dog
Red/Silver Fox
Nyctereutes procyonoides
Vulpes vulpes
OTHER MAMMALS
Bison Bison bison
Asian Elephant Elephas maximus
Capybara Hydrochaeris
hydrochaeris
FARMED IN
Canada, USSR,
Sweden
Australia, USA
Eurasia, Australasia,
N America
Eurasia, Australasia,
N America
Asia, Australasia
Australia, Mauritius,
Malaysia
China, Thailand,
Taiwan
UK, USA
China
China, N India
South Africa, Kenya,
Zimbabwe
South Africa,
Namibia, USA
South Africa, USA
Kenya, USA
Kenya, USA
Kenya
South Africa,
Namibia, USA
USA, Kenya,
Zimbabwe
South Africa, Kenya,
Zimbabwe, USA,
USSR
Europe, USA, USSR
N America, Canada,
Europe
USSR
Europe, USSR, China,
USA
Europe, N America,
Asia
Europe, USSR
Europe, China
N America, Europe,
USSR
N America
India, Laos, Myanmar,
Thailand
S America
391
WILD IN
N Europe, Asia, N
America
India, Sri Lanka,
Nepal
S Europe, N Africa,
Europe, USSR, N
America
Asia, USSR, Japan
Indonesia
SW Asia
China
SW China, N Viet
Nam
E Asia
Southern Africa
South Africa
Southern Africa
E Africa
E Africa
Southern Africa
Southern Africa
Africa
Africa
N America, Europe,
Asia
S America
USSR, China
Europe
Canada
S America
China
Americas, Europe,
Asia, Africa
N America
S and SE Asia
S America
Animal Use
Domesticated and semi-domesticated vertebrates (excluding
fishes) used in wildlife farming
PURPOSE
Meat, velvet, milk,
draught
Velvet, meat, medicinal
Velvet, meat
Velvet, meat, medicinal,
trophies
Meat, velvet
Velvet, meat, medicinal,
reintroduction
Velvet, meat, medicinal,
reintroduction
Restocking, medicinal
Musk
Musk
Trophies, meat
Trophies, meat
Trophies, meat
Trophy hunting
Trophy hunting
Trophies, meat
Trophies, meat
Trophy hunting
Trophies, meat, milk
Pelt
Pelt
Pelt
Pelt
Pelt
Pelt
Pelt
Pelt
Meat, trophies
Timber extraction
Meat, pelt
2. Uses and Values of Biodiversity
Table 26.18
NAME
Domesticated and semi-domesticated vertebrates (excluding
fishes) used in wildlife farming (continued)
LATIN NAME
OTHER MAMMALS (continued)
Musk Ox
Wild Boar
Cane/Grasscutter
Rat
Giant Rat
Vicufa
African Civet
BIRDS
Chukar Partridge
Red-legged
Partridge
Northern
Bobwhite/Quail
Emu
Common/Grey
Partridge
Common/Ring-
necked Pheasant
Ostrich
CROCODILIANS
American
Alligator
Common Caiman
Broad-nosed
Caiman
American
Crocodile
Australian
Freshwater
Crocodile
Morelet’s
Crocodile
Nile Crocodile
New Guinea
Crocodile
Estuarine
Crocodile
Cuban Crocodile
Siamese
Crocodile
OTHER REPTILES
*Green Turtle
*Hawksbill Turtle
Freshwater
Turtle
*Green Iguana
AMPHIBIANS
Frogs (various)
Ovibos moschatus
Sus scrofa
Thryonomys swinderianus
Cricetomys gambianus
Vicugna vicugna
Civettictis civetta
Alectoris chuka
Alectoris rufa
Colinus virginianus
Dromaius novaehollandiae
Perdix perdix
Phasianus colchicus
Struthio camelus
Alligator mississippiensis
Caiman crocodilus
Caiman latirostris
Crocodylus acutus
Crocodylus johnsoni
Crocodylus moreletii
Crocodylus niloticus
Crocodylus novaeguineae
Crocodylus porosus
Crocodylus rhombifer
Crocodylus siamensis
Chelonia mydas
Eretmochelys imbricata
Trionychidea
Iguana iguana
Rana spp. (etc.)
FARMED IN
N America
Europe, USA
W Africa
West Africa
Peru
Ethiopia
Worldwide
Worldwide
Worldwide
Australia
Worldwide
Worldwide
Southern Africa, USA
USA
E Asia, S America
Italy
Colombia, Cuba
Australia
Mexico
Africa, Brazil
Indonesia, Papua New
Guinea, Singapore
Asia, Australia
Cuba, Viet Nam
Thailand
Cayman Is, Réunion,
Suriname
Indonesia
Asia
Costa Rica
Asia
WILD IN
N America, USSR
Eurasia
West and Central
Africa
Caribbean
S America
Africa
S Eurasia
S Eurasia
N America
Australia
Eurasia
Eurasia
Southern Africa
USA
C and S America
S America
C and S America,
Florida
Australia
C America
Africa
Indonesia, Papua
New Guinea
E Asia, Australia
Cuba
E Asia
Asia
C and S America
Asia
PURPOSE
Meat, wool
Meat
Meat
Weed control
Wool
Musk
Meat, hunting
Meat, hunting
Meat, hunting, feathers
Meat, skin, feathers
Meat, hunting, feathers
Meat, hunting, feathers
Meat, skin, feathers
Skin, meat
Skin, meat
Skin, meat
Skin, meat
Skin, meat
Skin, meat
Skin, meat
Skin, meat
Skin, meat
Skin, meat
Skin, meat
Meat, shells, oil, leather
Shells
Pet trade, Meat
Meat, restocking
Meat (legs)
Source: Unpublished 1984 survey of wildlife ranching operations by WCMC (WTMU).
Note: Includes principal species only, grouped for convenience according to taxonomic group or by primary purpose. “Indicates semi-
domesticated species bred on a small or experimental scale
392
these environments but must also be adapted to their
commercial environment. Animal husbandry is the art or
science of reconciling domestic animals with these three
environments. Animal breeding is the art or science of
enhancing the positive economic response to husbandry, in
such a way that this enhancement is inherited. The basic
principles of animal husbandry are control over the
movements of the animals, the securing of supplies of feed
and water, and the management of reproduction so that
young are born at the right time of year.
Domestication and genetic variation
Domestic and domesticated animals with all their breeds and
types appear much more variable than their wild
progenitors. Domestication represents a genetic bottleneck,
meaning that the small sample of wild animals that is taken
into reproductive isolation may lack much of the genetic
variation of the wild population. However the great range
of breeds with all their different inherited characteristics
argues that either the genetic bottleneck was not particularly
narrow, i.e. that the total sample of wild animals was not
small, or that much genetic variation has arisen since
domestication.
Probably, many small new domesticate groups were set up,
most of which succumbed to the deleterious effects of
inbreeding. The minority that survived were groups which
thrived under these novel conditions, perhaps possessing
genetic material which predisposed them to cope with the
stresses of life with man (Kohane and Parsons, 1988), and
these groups then gave rise to all present domestic animals.
One attribute of a surviving inbred line or lines might be a
genetically determined tolerance of inbreeding (Templeton
and Read, 1984). Such tolerance could vary from breed to
breed, but it does seem clear that livestock in general show
declines in commercial productivity at similar levels of
inbreeding as those which zoo and wildlife managers or
laboratory animal specialists try not to exceed (Thomas,
1990; Roberts, 1982). In terms of biochemical
polymorphisms, domestic animals are at least as
heterozygous as wild populations, and often more so (Table
26.19).
Table 26.19 Mean heterozygosity in
selected vertebrate species
Mammals (in general) 0.041
Man 0.063
Cattle (3 breeds taurine, Belgium) 0.069 - 0.084
Pig (Belgium, Austria) 0.029 - 0.067
(feral herds in USA) 0.027 - 0.053
Wild boar (Italy, France, Austria) 0.021 - 0.031
Mouse 0.088
Domestic cat 0.066
Wild cat 0.042
White tailed deer 0.049 - 0.104
Moose 0 - 0.047
Red deer and Wapitu 0 - 0.060
Fallow deer (Britain, Italy) 0 - 0.006
Cheetah 0.013
Quail (domestic and wild 0.086 - 0.106
populations, Japan)
Mustelids (8 species) 0 - 0.060
Source: compiled from multiple sources.
393
Animal Use
Many breeds have been divided into strains which, while
specialised, are still closer to each other than to other
breeds (Hall, 1990). Originally, strains probably developed
as a simple result of herds and flocks tending to acquire
breeding stock from nearby areas. In 19th century Britain,
new breeds spread from their points of origin by slow,
steady diffusion (Walton, 1983, 1984). This pattern of local
interchange of breeding stock was described by Lush (1943)
as ideal for the adaptation of the breed as a whole to its
local environment.
General patterns of world livestock farming
As the human population of the world continues to grow,
the production of livestock will increase. Some of this
increase will come from further conversion of natural
habitat to agricultural use, while the rest will come from
intensification. Intensification means increased production
per livestock unit, or per hectare, and its pattern will vary
according to socio-economic conditions in the country
concerned.
Intensification will make the agroecosystem less diverse;
extensification will jeopardise natural and semi-natural
environments. The thrust of most development programmes
has been towards intensification; for example, provision of
deep wells in the Sahel region (aimed at extending the
grazing season and increasing the utilisation of a given
area). Perhaps because the developed world understands
intensification, which is market and science led,
development programmes are usually on these lines, and it
has generally been concluded that intensification is the only
practicable future course (Payne, 1986).
Patterns of cattle husbandry have been discussed by Meyn
(1984). Some 15% of Third World cattle are kept by
pastoralists (important in arid and semi-arid areas of Africa,
Middle East, parts of India and Pakistan, and central Asia).
About 30% are kept on ranches, which are mainly in Latin
America. The remainder are kept by smallholders. These
include a great variety of crop-livestock systems throughout
the tropics and sub-tropics, differentiated by altitude and
climate; and livestock farming without land as in Caribbean
feedlots and among Indian dairy farmers who supplement
roadside grazing with purchased fodder.
For other species, though there are census figures and
general qualitative descriptions of husbandry systems, there
are no global estimates of how the populations are
partitioned among the systems. Sheep tend to be kept in the
following general ways (Howe and Turner, 1984):
sedentary (sheep graze out from a home base, on crop
stubble, roadsides, waste land, steep hills); transhumant
(sheep move between summer and winter quarters, usually
highland and lowland respectively), and nomadic (no home
base, tending to move along well defined routes).
Generally, goats are kept in a similar range of systems.
The kinds of husbandry under which each species thrives
and yields a profit are determined by its biology. The goat
and the sheep are best suited to extensive systems, that is,
they can thrive on minimal husbandry. The pig has a
tremendous ability to revert to the feral state (in Australia
there are probably about 13.5 million feral pigs; Hone,
2. Uses and Values of Biodiversity
1990) and without fencing cannot be kept profitably in free
range. Chickens are too vulnerable to predators to be even
partly independent of man. Cattle need large amounts of
water to drink and large amounts of fodder. They, with
sheep and goats, are capable of ranging long distances in
search of food. Water buffalo need wallows in hot climates.
Cattle can be kept under tightly controlled conditions but
goats and sheep are less amenable to intensive rearing.
Cattle and water buffalo are the most versatile species to
market because they can be used for meat, hides, milk,
production of dung, and work. Further, cattle in rural
communities usually carry greater social prestige than other
species, except horses (sometimes) and camels
(occasionally). As a result of these factors, most livestock
development programmes worldwide have emphasised
cattle.
Animal production in developing countries
Animal production has a great deal to contribute to the
short- and long-term alleviation of individual and national
poverty. The special place animals hold in food supply
systems arises thus:
e animals can use wastes otherwise useless and can supply
traction and fertilizer
animals provide a form of low risk savings account
if milked, mammals can provide daily income
animal husbandry, being a year-round necessity, can
provide stable rural employment.
To be sustainable, systems should be based on locally
abundant feeds and human resources. The genetic potential
of the animals for production should be matched to the
resources available and this, typically, is best achieved by
the use of local breeds. In many developing countries,
animal production is not important at present and animal
products are bought in rather than produced locally.
Nevertheless, its wider adoption would help improve the
quality of life of many people. For instance, the humid zone
covers 19% of tropical Africa but has only 5% of the
domestic ruminant population; even though feed is plentiful
and there are many big cities, animal production has been
neglected partly for reasons of tradition (Armbruster and
Peters, in press).
Provided enough resources are allocated, probably any
breed of livestock can survive and produce in any country,
though the substantial recurring expenditure on imported
feed, veterinary care and housing may mean that unless
subsidies are forthcoming the enterprise would not be
profitable. Intensive industrial farming systems such as
those based on Holstein-Friesian cattle, Large
White-Landrace pigs, and hybrid fowl, need not directly
supplant local breeds. These enterprises are not sustainable
in less-developed countries, however, and many regard
them as entirely inappropriate subjects of aid funding. Such
aid programmes have been numerous in South America,
where large numbers of pure-bred North American and
European dairy cattle have been sent (120,000 to Venezuela
alone in the period 1983-88). Mortality rates have been
extremely high and the system is far from being sustainable.
394
However, it is clear (Vaccaro, 1990) that an element of
crossbreeding with local cattle that are adapted to the
environment greatly improves survival without excessive
penalties on milk yield.
One advantage of the continued existence of local breeds of
livestock therefore is that they provide genetic material to
enable imported breeds themselves to become locally
adapted, and thus to help rescue schemes which were put in
place without adequate planning.
Exploiting genetic diversity
The biological diversity represented by a multiplicity of
different breeds enables productive agriculture to be carried
out in a wider range of environments than would be the
case if there were genetic uniformity. The local adaptations
of breeds can reduce dependence on veterinary care.
Breed diversity also permits more rapid genetic progress to
be made. It is always quicker to develop livestock by
importing genes from outside than by selecting within a
breed. One breed can act as a source of genetic material for
another. This reservoir began to be tapped as husbandry
developed and market requirements changed, leading
farmers and breeders to look elsewhere for breeds that
could be mated with their own stock to produce more
remunerative animals. For example, in the 18th century,
Merino sheep, Chinese pigs and dairy cattle from the Low
Countries (Hall and Clutton-Brock, 1989) were all imported
and crossed with local British types to confer on them
fleece quality, pork quality, and milk yield, respectively.
Sometimes, new genetic mutations manifest themselves in
flocks and herds and these can act as the foundation of a
new breed. The best known in recent years has been the
Booroola gene found in certain Merino sheep, which
enhances ovulation rate (Bindon and Piper, 1986). Another
instance led not only to the foundation of a new breed but
also a new industry. In 1931 a mutation in a New Zealand
Romney sheep resulted in a ram lamb with a very hairy
fleece. The gene for hairiness, when homozygous, resulted
in a fleece 65% by weight of hair, 35% wool. This mix
turned out to be ideal for carpet manufacture, which was
previously not economic in New Zealand. The new breed,
the Drysdale, has attained some importance (Nicholas,
1987).
During the 19th century British livestock breeds were
exported to be crossed with local types all over the world
(Hall and Clutton-Brock, 1989); exports dropped sharply
during the first half of the 20th century and more attention
was then paid to local stocks. Today there is a great deal of
pressure on tropical countries to accept North American
dairy cattle; hundreds of thousands of Holsteins in
particular have been exported, notably to Latin America
(Vaccaro, 1990). However more and more advisors are
maintaining that purebred temperate zone breeds like these
are not appropriate in such areas and more attention should
again be paid to local breeds (McDowell, 1985; Bondoc et
al., 1989; Vaccaro, 1990; Wilkins, 1991).
The crossing of breeds can be conducted according to the
following systems.
Breed replacement
This took place several times on the plains of North and
South America. Range cattle of Spanish descent were run
as vast semi-feral herds primarily for hide production until
the development of railways, refrigerated ships and cold
stores led to expansion of the beef market. It then became
worthwhile crossing the range cattle with British breed
bulls, first Shorthorns, then Aberdeen Angus and
Herefords. More recently still, arid lands have been made
into ranching areas by the use of drought adapted cattle
such as the Santa Gertrudis, developed by adding genes of
zebu bulls (Sanders, 1980). Contemporary North American
Tange cattle and those of the pampas of South America
probably include in their genetic makeup only a tiny
proportion of Spanish genes, but these genes, with those of
the later imports, were the material on which a combination
of natural and artificial selection has acted to produce
locally-adapted animals. Only a very few cattle considered
to be Texas Longhorns (the original Spanish stock of the
south-west) and Florida Scrub (that of the south-east)
survive and these are the subject of conservation efforts
(Simmons-Christie, 1984; Olson, 1987).
Formation of a synthetic breed
This can result from crossing two or more breeds and then
selecting from the crossbred stock. Examples include the
Jamaica Hope dairy breed (McDowell, 1985), stabilised at
80% Jersey, 15% Sahiwal (one of the very few breeds of
zebu dairy cattle), and 5% Holstein. There has also been
much crossbreeding of European and North American dairy
cattle (of the taurus group) with local zebu breeds, in South
America and India notably, mainly for milk production
(Cunningham, 1989). Typically, age at first calving and
calving interval are reduced, and first lactation milk
production is increased up to 50% in step with the
increasing proportion of introduced genes. If that proportion
is exceeded, calving interval tends to be longer and milk
performance not much improved.
Stabilised crossbreeding
In this system, breeds are bred pure but the progeny
crossed. This combines in the offspring the merits of both
parents. In some such systems the offspring are superior,
with respect to traits of value, than the parental mean; that
is, they exhibit hybrid vigour (heterosis). The standard
technique for exploiting genetic distinctiveness has been to
make use of the additive or heterotic effects that can arise
when distinct breeds are crossed (Hall, 1990).
Strategies for genetic improvement of livestock
Use of locally existing genotypes
Advantages of this course are that such livestock may well
be adequate and able to respond sufficiently to
improvements in the system. Over many generations they
will have acquired the ability to perform locally appropriate
and multiple functions. They will probably have resistance
to local diseases. Breeding stock would be locally available
and their purchase would create cash flow and contribute to
the confidence of those who are particularly competent
breeders. The disadvantages are that it is not as glamorous
an approach as the importation of new genotypes and is
perhaps less likely to attract aid funding as it does not
involve heavy expenditure on imports from donor countries.
395
Animal Use
Local breeds can be improved by selection without the
admixture of imported genetic material, but it is hard to
predict whether the results would justify the investment.
This is because heritabilities of commercial traits are
difficult to estimate (see Table 26.20), the possible selection
intensities are likely to be low, and the programme depends
critically on the collection and analysis of records. However
the scheme most likely to work is a nucleus breeding
scheme, whereby participating breeders contribute their best
females to a central unit and are entitled to purchase stud
males from the unit. Such a scheme apparently operates in
Libya, with the Libyan Barbary fat-tailed sheep. Howe and
Turner (1984) reported that since 1978 20,000 ewes in
small flocks had been screened and a nucleus of 2,000 ewes
established and subjected to selection. Rams from this
nucleus flock are distributed back to the small flocks.
Table 26.20 Heritability of various traits
in animals
CATTLE
Birth weight
zebu, tropics 0.38
taurine, temperate 0.45
Weaning weight
zebu, tropics 0.29
taurine, temperate 0.26
Heat tolerance (zebu x, Australia) 0.44
Tick burden (zebu x, Australia) 0.39
Worm egg count (zebu x, Australia) 0.12 -0.25
Milk and component yield (Holsteins, USA) 0.25
Body size traits (Holsteins, USA) 0.40
Milk composition traits (Holsteins, USA) 0.55
Disease susceptibility
Mastitis (Holsteins, USA) 0.01 - 0.07
Feet and legs (Holsteins, USA) 0.10
Sum of all diseases (Holsteins, USA) 0.02 - 0.06
Milking behaviour (Holsteins, Canada) 0.12 - 0.16
Ease of handling 0.12
Aggressiveness at feeding 0.11
Calving interval (zebu, Brazil) 0.23 - 0.86
Lifetime number of calves reared 0.03
SHEEP
Clean fleece weight 0.45
Fibre diameter 0.12 - 0.50
Staple length 0.30 - 0.60
Embryonic mortality (Romanov, France) 0.09
Litter size (Romanov, France) 0.02
Ewe fertility 0.00 - 0.17
Lambs born (per ewe lambing) 0.04 -0.15
OTHERS
Mohair and cashmere (goats)
Fibre length 0.70
Fibre diameter 0.12 - 0.40
Energetic efficiency (broiler fowl) 0.30 - 0.40
Liability to myxomatosis (rabbits) 0.35
Stature (humans) 0.51
Source: compiled from multiple sources.
Notes: The higher the heritability (range 0-1 .0) the greater the response
to selection in the environment.
Replacement by imported genotypes
The importation of Spanish cattle to the Americas, of
Merino sheep to Australia, and the rapid contemporary
2. Uses and Values of Biodiversity
spread of the Holstein-Friesian are examples of this
process. However, whilst industrial farming is generally not
appropriate in developing countries, the possible role of
imported breeds in traditional husbandry bears examination
because of the basic fact that development of a breed by the
introduction of genetic material from other breeds is much
more rapid than the development of a breed by selection.
One result of most published studies on local breeds has
been to show that they already possess the genotype
enabling them to respond to improved husbandry and this
could make the importation of genetic material unnecessary.
Steinbach (1986) established an experimental herd of goats
on a research station in Tunisia and compared the local
nondescript breed with the Boer, a breed specially
developed for meat production, and the Alpine, Saanen and
Poitou (European dairy breeds). He found the local breed
to be the most profitable, responding very well to improved
husbandry and incurring the least veterinary expense.
Similarly, Nguni cattle (Scholtz, 1988) from the east coast
of southern Africa perform comparably to improved breeds
in controlled trials under good husbandry.
Importing exotic stock into developing countries is generally
a high cost, high risk strategy, which is unlikely to solve
the problems of the majority of farmers. It is advantageous
for donor countries because it provides continuing profits
for breeders and for veterinary products and services.
Disadvantages have come to light as a result of practical
experience; very many introductions, particularly of dairy
cattle, have failed and others are kept going only by
massive and continuing importations of replacement
females.
Supplementation with imported genotypes
The introduction of Indian humped (zebu) cattle to the
Americas (Sanders, 1980), mostly over the last 100 years,
illustrates this process. Crossbreeding among the imports,
with little if any contribution from European cattle, led to
the Brahman, Indu-Brazil, Gir, Guzera and others;
crossbreeding in Texas with pre-existing British type cattle
led to the Santa Gertrudis which is 5/8 Shorthorn and 3/8
Brahman.
Recent examples of breeds being imported to add genetic
material to local breeds include the highly prolific Meishan
pig, one of the Taihu breed group of China, now being
widely used in breeding programmes in Europe (Sellier and
Legault, 1986), also highly prolific, and the Sahiwal dairy
breed of Pakistan, imported to Australia from 1960 to
confer tick resistance on Friesian cattle (Turton, 1985). The
Finnish Landrace sheep has been used to develop new
breeds such as the Cambridge (Owen and ap Dewi, 1988)
and in crossbreeding schemes, most notably with the Dorset
Horn, to produce ewes which are mated with rams like the
Suffolk to produce meat lambs. None of these breeds is
considered rare in its native country, but it is quite possible
that rarer and less well known breeds may well exist there
which themselves may possess useful genes.
The most dramatic livestock development of the last 200
years, the emergence of range cattle husbandry in North
and South America, arose through progressive
crossbreeding of imported British breeds with the Texas
396
Longhorn and other Criollo breeds descended from those
brought from Spain soon after Columbus. Here, a slow
process of natural selection led the cattle, which with each
successive generation resembled more and more their
purebred British ancestors, to retain the locally adapted
genes of their Criollo ancestors. This is the process of
upgrading. If inseminations of native cows are only by
imported bulls, the average percentage of the genotype of
the progeny that is of imported type will increase from
generation to generation in the progression 50%, 75%,
87.5%, 93.75%. In principle, the small percentage of native
genotype remaining comprises, by natural selection, the
genes adapting the animal to the local environment.
Most tropical breeds of cattle have only low milk yields
(Turton, 1985) but if crossed with temperate zone dairy
breeds, yields of the progeny are at least double those of
the local breed. The practicalities of a crossbreeding scheme
that maintains the proportion of temperate zone blood in the
milking cows at 50% (which has generally been found to be
sufficient for maintaining high milk yield without
jeopardising local adaptation) are complicated, and it seems
essential that a continued input of genetic material from the
temperate zone breed is necessary. An alternative would, in
principle, be to create a new breed, by mating among the
first crossbred generation and then selecting, and this has
been tried, but the synthetic dairy breeds thus created in
tropical countries have not generally been very successful.
Replacement of local stock by nearby breeds
Sometimes local breeds may be replaced by supposedly
more profitable breeds from the same or neighbouring
countries. In Nigeria, for example, West African Shorthorn
or Muturu cattle, a dwarf trypanotolerant breed of the
coastal and central: zones (adult body weight about 200kg)
is under threat of replacement by other West African
breeds, though as the breed is still numerous this is a
long-term threat. These cattle are kept under a form of
communal ownership in villages where the main interest is
in crop growing. Numbers suffered greatly in the civil war
of the late 1960s and have not recovered, there being little
local interest in their husbandry for profit. Schemes aimed
at promoting cattle raising in these areas are based on the
NDama, another trypanotolerant breed, mainly from
Senegal. In the central zone, tsetse fly eradication and a
preference by traders for larger bodied cattle mean that the
White Fulani or Bunaji, a humped apparently
trypanosomiasis-sensitive breed, has been replacing the
West African Shorthorn. It is ‘also possible, though data are
lacking, that the Kuri (a large bodied humpless breed with
giant bulbous horns), kept in the Lake Chad area by
sedentary communities, could be under pressure as a
consequence of fighting in Chad, the spread of cultivation
around the Lake, and perhaps by pressure from the Red
Bororo cattle kept by migratory pastoralists.
Use of wild relatives of domestic stock
There are many examples where plant collecting expeditions
to areas of diversity for domesticated plants and their wild
relatives have brought back genetic material of great value
for crossing with cultivars (e.g. to improve hardiness and
disease resistance). The use of interbreeding with wild
animals to improve domestic livestock is much more
uncommon, possibly because of the smaller number of
species and the extreme rarity of most wild relations of
domestic species. However, there is some potential for this:
for instance, some of the impetus for tracking down the
remaining Kouprey Bos sauvelii is the belief that they may
possess natural immunity to various diseases which could be
harnessed by cross-breeding.
Rare or threatened breeds
Pursuit of higher production targets, the commercial success
of particular breed promoters, and, in developed countries,
changes in consumer preferences have led to livestock
development activities becoming concentrated in few breeds
and breed groups. The corollary of this is that more breeds
are declining in importance, many have been lost and the
survival of many others is in considerable doubt. Concern
for rare breeds has been most marked in north temperate
countries with a history of specialised livestock production,
but it is becoming increasingly evident that declining breeds
in less developed countries can represent genetic resources
of great significance. Here, it seems likely that local
varieties distinct enough to be defined as breeds had
European criteria been applied may already have been lost.
The lack of inventories and of status reports for local
breeds in developing countries is cause for concern, as is
the lack of support for local breeds in development
programmes.
The most authoritative world list of animal breeds (Mason,
1988), lists a total of 3,237 extant breeds of ass, buffalo,
cattle, goat, horse, pig and sheep. The number of such
breeds in each country with native breeds is shown in Table
26.21. Data for certain countries are shown graphically in
Fig. 26.9; the countries have been selected to illustrate
general global patterns of breed richness.
Some 474 of extant breeds can be regarded as rare (Hall, in
press). A further 617 have become extinct since 1892;
numbers of extinct breeds in each country are given in
Table 26.22, and data for selected countries are shown in
Fig. 26.10.
Overall breed numbers (extinct, rare and non-rare) in each
continent are shown in Table 26.23.
There is likely to be significant bias in these data,
particularly with regard to extinct breeds. For example, of
the 1,259 cattle breeds listed by Mason .(1988), 242 are
indicated as extinct, of which 200 were in Europe and the
former USSR; only 20 were in Africa and two in India.
Because breeds tend to be less formally structured and not
well documented in developing countries, genetic variation
may not be adequately represented by current breed
nomenclature. However, it may be that in developed
countries where human populations are high, rates of breed
development have also been high, in response to
commercial and aesthetic demands. Breed turnover, as
measured by numbers of extinct breeds, would be expected
to be high in such circumstances. Whatever the explanation
may be, present data show that the great majority (83%) of
known breed losses occurred in Europe and the former
USSR.
397
Animal Use
Reasons to conserve breeds
Threatened breeds ought to be conserved for the following
economic reasons:
e they may possess adaptations to local conditions,
@ they may possess adaptations which can be exploited in
other geographical areas or farming systems.
One of the great advantages of having access to a diversity
of breeds is that in several, breed characteristics exist which
are governed by single genes. In principle, single
favourable genes could be transferred from one breed to
another (Davis and Hinch, 1985). In practice, the major
gene may owe at least some of its efficacy to its genetic
background, and in a recipient breed the background may
be different and unpredictable. Even detecting an animal
that carries the Booroola gene (see above) is difficult
(Haley, 1991). Genetic probes can be used to identify
genotypes; it has proved possible to treat spermatozoa with
such probes to identify which individual bulls carry a
certain gene (coding for kappa-casein) which improves the
suitability of milk for cheesemaking (Medrano and
Aguilar-Cordova, 1990). In California, the Jersey breed has
a far higher frequency (88%) of cows homozygous for this
gene than does the Holstein.
Rare breeds in protected areas
Several protected areas provide a home for notable feral
populations; Chillingham Park in northern England has been
the home of the Chillingham white cattle possibly since the
13th century (Hall, 1989a,b), and the St Kilda islands off
north-west Scotland are a refuge for the Soay sheep (Jewell
et al., 1974). In New Zealand, a reserve for feral sheep
was established on Pitt Island (Rudge, 1983).
There are some countries where protection has been applied
to endangered breeds through areas in national parks being
set aside for them. These include Ireland (Muckross
National Park: Kerry cattle; O’hUigin and Cunningham,
1990), Hungary (Hortobagy National Park: Hungarian Grey
cattle, Mangalica pigs, Racka sheep; Henson, 1983), Poland
(Roztocze National Park: Konik pony; Sasimowski and
Slomiany, 1987), Swaziland (Mkhaya Farm: Nguni cattle;
Setshwaelo, 1990).
Information requirements
The first step in organising conservation is to compile an
inventory and to decide on priorities. Examples of
inventories are cited by Hall (1990). Worldwide, FAO is
organising a global data bank (Maijala, 1990) while the
European Association for Animal Production has published
a list of endangered populations in Europe, to the number
of 241 (Maijala et al., 1984).
Many breeds of livestock are promoted by breed societies.
In the British tradition of pedigree breeding, which has been
adopted in very many other countries, the breed societies
each operate a register of breeding stock, known as a stud
(equine), flock (sheep), or herd (cattle, goat, pig) book.
Such societies are almost entirely lacking in the developing
world.
2. Uses and Values of Biodiversity
Table 26.21 Numbers of extant breeds in each country with native breeds
ASS BUFFALO CATTLE
ASIA
=
>
Afghanistan
Bahrain
Bangladesh
Bhutan
Burma
Cambodia
China 6
Cyprus 1
India 1 6
Indonesia 1
Iran 4 1
1
1
=
NH fF
Re
=
(1)
oa
RH] NOTWMAAAWAIN-HN
Iraq
Israel 1
Japan
Jordan 1
Korea N and S
Laos 1
Lebanon
Malaysia 1
Mongolia
Nepal 3
Oman
Pakistan 4
Philippines 2
Saudi Arabia
Sri Lanka
Syria 2
Taiwan
Thailand
Turkey
Vietnam
Yemen 2
USSR (former)
(1)
PHN HD
_
a=
(1)
—=---) 0
=
PUOWOnH—- as
15 1 58
EUROPE
Albania 1 1
Austria
Belgium
Bulgaria 1
Czechoslovakia
Denmark
Faeroe Islands
Finland
France 2 (1)
Germany
Greece 1
Hungary ‘
Iceland
Ireland
Italy 8 (6) 1
Malta 1
Netherlands 10
Norway 7
Poland 5
Portugal 18
Romania 2 9
Spain 7 (3) 37
Sweden 5
Switzerland 6
UK 42
Yugoslavia 1 19
NORTH AND CENTRAL AMERICA
Central America (gen)
Bahamas
Barbados
Belize
Canada
Costa Rica
Cuba
Dominican Rep.
i“)
HH ANADANT-AAIainigns
wR
wo
©
oanw o
(1)
(2)
(3)
(2)
(9)
398
GOAT
>
w
OwWAN AHN RHF HAH]
wo2=Ss-aann==
o- fn
(2)
(4)
(1)
(6)
(1)
(1)
(1)
(11)
(1)
(2)
(1)
(3)
(3)
HORSE
5
1
1
1
16
7 (1)
9
12° «(1)
1
8 (8)
2
1
5
4
7 (3)
4
1
1
9 (1)
1
9
60 (23)
3
5 (1)
4
11
7 (1)
3 (2)
1
1 (1)
19 (6)
20 (10)
3
10 (1)
2
4 (1)
11 (9)
5 (2)
4 (1)
21 (2)
5
8
8 (2)
5 (1)
1
20 = (7)
7 (1)
5 (2)
PIG
103
34
wR
NNO
“
—-= =
CONN O-UM— Ww
=
(1)
(1)
(2)
(1)
(1)
(1)
(7)
(4)
(6)
(2)
(1)
(1)
(4)
(8)
(2)
(1)
SHEEP
Shin ior
133
wo
Ore =| NON FD
5
aned
(1)
(11)
(2)
(1)
(13)
(2)
(13)
(6)
(7)
(2)
(24)
(1)
(10)
(2)
(2)
(4)
(4)
(17)
(5)
(1)
Animal Use
Table 26.21 Numbers of extant breeds in each country with native breeds (cont.)
ASS BUFFALO CATTLE GOAT HORSE PIG SHEEP
NORTH AND CENTRAL AMERICA (continued)
Guadeloupe 1
Guatemala (1)
Honduras 1
Jamaica 1
Mexico
Nicaragua
Puerto Rico
United States 6 4
SOUTH AMERICA
South America (gen) 1 1
Argentina 5
Bolivia 4
Brazil 5 (1) 2 26 = (4) 8
Chile 1
Colombia 9
Ecuador 2
1
1
=
(1) 1 2 4 (1) 4
o--WFf
(6) 10. (4) 35 (7) 10 (4) 32 (6)
13 7
== 0-0
Paraguay
Peru
Uruguay 1
Venezuela 5 1 1
OCEANIA
oa
=
(1)
RH
Oceania (gen) 1
Australia 20 2 2 23° (2)
Fiji 1
Guam 1
Hawaii 2 (1) 1
New Zealand
N Marianas 1
Papua New Guinea 1
AFRICA
Algeria 1
Angola
Benin
Botswana
Burkina Faso
Cameroon
Chad
Cote d'Ivoire
Egypt 3 4
Ethiopia 2
Gambia
Ghana
Guinea
Guinea-Bissau
Kenya 1
Liberia
Libya 1
Madagascar
Malawi
Mali
Mauritania
Morocco 1
Mozambique
Namibia
Niger
Nigeria
Senegal
Seychelles 1
Sierra Leone 1
Somalia 1
South Africa 11
Sudan 4 20 1
Tanzania 2 9 (1)
Togo 4
Tunisia 1 4
Uganda 12
Zaire 8
Zambia
Zimbabwe 4 3
=
oa
=
tS
=
a
(3)
(2)
=
(4)
QO-p2 2 =
w
On---f0
=
(1)
=
(1) 2 1
(1)
=
MD-TAMANWOHHDH FH WEWHWBHANHDANHNHNA UND
==]230.2-2 2-4
(1)
pS
OON=-=0NnO02-26h 20-22-06
(1)
=
-~O©W -$NWONN—
(2)
=2=nN0-22+a 02
O-O-nNODTN =
Source: data from Mason, 1988; analysis by S.J.G. Hall (1992. Livestock breeds and their conservation. 1. World distribution. in prep.)
Note: principal numbers are estimates of extant breeds, numbers in parentheses indicate those classed as rare by Hall (included in main figure).
399
2. Uses and Values of Biodiversity
Figure 26.9 Numbers of living breeds of livestock in selected countries
OO
ISNERSRASEASY
XK KK KKK XX
MIP‘GeWpsaq\|J\PIoJMeID
ee
Table 26.22 All countries: numbers of breeds extinct since 1892
CATTLE
ASS
ASIA
China
Hong Kong
India
Japan
Pakistan
Philippines
Taiwan
Turkey
USSR (former)
EUROPE
Austria
Belgium
Bulgaria
Czechoslovakia
Denmark
France
Germany
Greece
Hungary
Ireland
Italy 4
Netherlands
Norway
Poland
Portugal
Romania
Spain 1
Sweden
Switzerland
United Kingdom
Yugoslavia
NORTH AND CENTRAL AMERICA
Canada
United States
SOUTH AMERICA
Brazil
Chile
Uruguay
Venezuela
OCEANIA
Australia
New Zealand
AFRICA
Algeria
Benin
Cameroon
Gambia
Lesotho
Malawi
Nigeria
Rwanda
South Africa
Tanzania
Zimbabwe
TOTAL 5
22
=
=)
ROH HH Wu
LS in OL!
228
GOAT
26
HORSE
20
90
PIG
21
18
24
17
126
Animal Use
SHEEP
31
nyo
142
Source: data from Mason, 1988; analysis by S.J.G. Hall (1992. Livestock breeds and their conservation. 1. World distribution. in prep.)
Note: Grand total worldwide = 617.
401
2. Uses and Values of Biodiversity
Figure 26.10 Numbers of extinct breeds of livestock in selected countries
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403
2. Uses and Values of Biodiversity
Costs and benefits of conserving breeds
A breed can be conserved (a stock maintained which
continues to represent the foundation stock without too
much genetic drift or inbreeding) for surprisingly small cost
compared with the possible economic benefits. Either a live
breeding stock can be maintained, or semen or embryos
preserved, or all methods can be used. For semen and
embryos, the genetic variability in a typical breed would be
adequately represented by collection from 25 males, or by
25 embryos each from 25 donors (Smith, 1984). Embryo
storage is not yet possible for chickens and pigs, and is not
yet fully developed for equines (Guay and Poitras, 1989;
Heyman and Vincent, 1988).
Live breeding stocks are much more expensive to maintain,
as any farm that keeps a conservation unit of a
non-commercial breed is losing the opportunity to keep a
profitable breed. If it is decided to keep a live conservation
population, its size is best defined by what rates of
increment of inbreeding and of fixation of genes through
random genetic drift are permissible. An effective
population size of 30 seems appropriate (Smith, 1984). The
rates should be expressed per year not per generation and
recalculation yields the minimum sizes of population of each
species necessary to keep annual increment of inbreeding at
below 0.2%. These population sizes are surprisingly small,
provided appropriate sex ratios are chosen.
Potential benefits of livestock conservation are very great
(Smith, 1984). If a 1% gain in economic efficiency arises
in a livestock industry through the use of a conserved
breed, this benefit will exceed the cost by between 33 and
190 times. Even though the cost/benefit ratio of livestock
conservation is so favourable, in absolute terms the amounts
required are still large. In Europe alone, there are 241
breeds and strains that appear to justify conservation
(Maijala et al., 1984).
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K.H. (Eds), Neotropical Wildlife Use and Conservation. University
of Chicago Press, USA. Pp.6-23.
Roberts, R.C. 1982. Lessons to be drawn from selection experiments
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Sheep and Beef Cattle Breeding, Vol. 1. Dunsmore Press,
Palmerston North, New Zealand. Pp.253-259.
Ruddle, K. 1973. The human use of insects: examples from the
Yukpa. Biotropica 5(2):94-101.
Rudge, M.R. 1983. A reserve for feral sheep on Pitt Island, Chatham
Group, New Zealand. New Zealand Journal of Zoology,
10:349-63.
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The Food Insects Newsletter (TFIN):
1) Volume II, No.3 pp.2 and 10
2) Volume III, No.2 pp.1, 3, 4 and 6
3) Volume III, No.3 p.1
4) Volume IV, No.2 pp.3-4
5) Volume IV, No.2 p.8
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1982. What's Wildlife
2. Uses and Values of Biodiversity
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Section on domestic livestock abridged from a consultancy
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Mammalian Ecology and Reproduction, University of
Cambridge), with additional tables extracted by permission
from work in preparation by the same author.
27. BIODIVERSITY AND ECONOMICS
A growing literature in applied economics is demonstrating
that techniques are available for obtaining concrete
estimates of the value of many different facets of the
environment, including the more intangible aspects of
environmental quality, such as clean water, clean air and
better views. These methods can be applied to biodiversity,
but are subject to major limitations and problems of
interpretation. One of the major difficulties is that they are
based on the premise that value is determined by human
willingness to pay. The range of human values can be very
broad and consequently difficult to measure: many people
are willing to pay for qualities that are seemingly unselfish,
by placing, for example, an ‘existence value’ on certain
natural resources that they will never personally see or
experience.
The major problem, however, involved in the application of
these methods to biodiversity is defining exactly what is
meant by biodiversity, a notoriously intractable question.
In this regard, the distinction between valuing biological
resources and valuing biological diversity (i.e. the range of
variation in biological resources, whether measured
quantitatively or qualitatively) is an important one and leads
to two different types of question: in the first instance, a
gross estimate of the value of biological resources in a
particular geographic locale is sought; in the second,
attempts are made to trace the impact of changes in
diversity on economic values.
VALUING THE ENVIRONMENT
The total economic value of an environmental resource may
be broken down into a range of use and non-use values.
The direct use of ecosystem outputs in non-consumptive,
consumptive or productive activities is the impact that is
most commonly measured in valuation exercises. Included
as direct uses would be the harvesting of wild species for
use as food, fuel, shelter or medicine. Other activities such
as ecotourism involve a direct ‘transaction’ between people
and biological resources and fall into this category of direct
use values. Some direct uses of biological resources such as
commercial logging, agriculture or fisheries generate
products which are exchanged in the marketplace, while the
products of others such as subsistence hunting and gathering
go largely unmarketed. In the latter case, although these
non-marketed resources have no financial value (cash price
in exchange) they do have economic value as they are of
importance to society.
Biological resources may also make indirect contributions
to the welfare of society. Environmental functions support
economic activity by recycling important elements such as
carbon, oxygen and nitrogen and by acting as a buffer
against excessive variations in weather, climate and other
natural events outside the control of human beings.
Economists are increasingly attempting to place values on
these indirect use values. Since indirect use values do not
enter directly into human preferences and are often widely
available, their value is not often recognised and
incorporated into development decisions. As natural habitat
declines these ecological processes become scarcer, their
economic value grows and eventually mechanisms are
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designed for ‘marketing’ these services (note the increasing
trend towards user charges for water supply and waste
disposal in developed countries).
In addition to direct and indirect use values, biological
resources may have option and existence (non-use) values.
Option values are associated with the future use of a
resource and future flows of information regarding the use
of resources. Risk-aversion dictates that societies should be
willing to pay an additional sum above and beyond what a
future use value of a biological resource is worth in order
to guarantee future access. If this is the case, there is an
‘extra’ value that can be placed alongside the use values of
the resource.
Finally, there may be non-use or existence values associated
with a resource. These are benefits derived by an individual
from the mere knowledge that the resource exists. For
example, people who donate money to a conservation
organisation with no expectation of ever visiting the habitats
or hunting the species which the organisation aims to
conserve must be deriving some satisfaction that is simply
a result of the continued survival of the species or habitat.
To sum up the different types of values that make up the
total economic value of biological resources, Table 27.10
illustrates how the goods and services produced by a
tropical forest fit into this ‘taxonomy’ of values.
A given habitat or species may have many different use and
non-use values. Ancient redwoods may have ecotourism
value, timber value, contribute to watershed protection and
carbon storage, and have significant existence values.
Because a number of values may be involved and since
some techniques are better than others for measuring
different types of values, any comprehensive valuation
exercise of a particular habitat or even one species may
involve the application of a range of valuation techniques.
Care must be taken, however, to avoid simply adding the
resulting values to each other to obtain a total economic
value. Trade-offs between values and double-counting of
benefits may occur, making simple summation of the
outcomes of separate analyses of different values potentially
very misleading. Despite these difficulties the techniques
reviewed below provide useful methods for quantifying the
benefits of environmental resources.
Changes in productivity approach
The changes in productivity approach relies on an
understanding of underlying ecological relationships to
derive a model indicating how changes in the supply of an
environmental resource results in changes to the economic
value of production. This technique can be used to
investigate improvements or damage to environmental
quality. For instance, soil fertility has a direct impact on
agricultural productivity. Soil degradation will raise crop
production costs for a certain level of output. Resulting
changes in quantities and prices will cause the benefits
received by consumers and producers to change. Comparing
initial levels of surplus with the resultant levels provides a
way of estimating the value of changes in supply of the
environmental resource or quality. This technique is
2. Uses and Values of Biodiversity
particularly relevant to basic resource issues in developing
countries where a large proportion of economic production
comes from agriculture, fisheries, forests, etc. The
production function technique is a natural complement to
cost-benefit analyses of projects that require estimation of
the economic effects of changes in resource availability.
The Nepal Hill Forest Development Project provides a
simple example of using the changes in productivity method
to value improvements in environmental quality for
incorporation in cost-benefit analysis. As reported by Dixon
et al. (1988) the project involved introducing systematic
hill-forest development into 38,500ha devoted to a mixture
of land uses in the vicinity of Kathmandu and Pokhara. The
objectives of the project were to reduce soil erosion,
increase the productivity of different land uses in the
watershed and provide sustainable flows of fuelwood and
fodder, amongst other resources. The benefits from
reductions in soil erosion were not quantified, but
improvements in the physical yields of milk, fertilizer and
fuelwood were calculated for the four land types: grazing
land, pasture, unmanaged scrubland, and unmanaged forest.
Using readily available market values the project values for
milk and fertilizer production were calculated.
Three different methods were used in calculating the unit
value for increased fuelwood supplies. A direct market
value approach used the economic price of fuelwood (minus
transport costs) in Pokharaand Kathmandu (280 rupees/m’).
As the production from the project would increase
Kathmandu’s fuelwood supply by 20% and because the two
markets were considered small and isolated, two additional
techniques were utilised. Cattle dung is the closest available
substitute for fuelwood in rural areas. The economic price
of this fuelwood substitute was estimated to be 65 Rs/m’
based on the marginal loss of foodgrains that would occur
if dung was diverted away from its role as a fertilizer. A
final approach involved valuing fuelwood in terms of the
opportunity cost of labour diverted from other employment
by the need to gather fuelwood. The opportunity cost
approach yielded a value of 83 Rs/m? for fuelwood. The
correct value to use in calculating fuelwood production is
the lowest value - this case that derived from indirect
substitution. The total production values for the different
land types were aggregated and compared with the benefits
from allowing continued land and forest degradation.
Contingent Valuation Method (CVM)
The objective of any valuation exercise is to determine
people’s preferences for environmental quality. What this
entails is discovering what people are willing to pay (WTP)
for increments in environmental quality or what they are
willing to accept (WTA) in compensation for forgoing such
benefits. While there is little theoretical reason for
suspecting that WIP and WTA should be different,
empirical research has revealed that measurements of WTA
often exceed those of WTP. In this discussion, WTP is
used as representative of demand for environmental quality.
By undertaking surveys or administering questionnaires it
is possible to elicit people’s WTP (or WTA) for
environmental goods or services directly. By creating a
hypothetical market situation the researcher can use the
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respondent’s replies to place values on items that are
usually not marketed. The valuation is ‘contingent’ because
the values derived from CVM depend on individual
perceptions of a host of background factors that influence
the market being surveyed. A poorly designed and
implemented survey will produce answers that bear little
resemblance to the population’s true WTP. It is precisely
because there is so much room for difference between
consumer intentions as expressed on a questionnaire and
consumer preferences as revealed in the marketplace that
CVM results are often considered unreliable.
The literature has focused on overcoming the many sources
of bias in CVM studies. Bias is any element in the study
that consistently skews results in one direction, thereby
leading survey results away from the true WTP of the
population. Biases may arise from the way the sample is
selected, the effect of the survey design or implementation
on the responses gathered from respondents, or when the
respondent misunderstands the nature of the contingent
market. Resolving these difficulties involves careful design
and testing of questionnaires, competent survey
administration and a number of econometric tests for
remaining sources of bias.
The use of CVM for valuing environmental resources is
largely a North American and, to some degree, European
tradition with very little work conducted in developing
countries. A recent study in Brazil, however, indicates that
results of CVM studies are credible even in rural areas in
developing countries when respondents are well-informed
about the resource in question. The study demonstrated how
CVM surveys of actual and hypothetical water-use practices
can provide estimates of WTP for access to clean water that
vary according to household socioeconomic characteristics,
and qualitative differences in water supply and delivery
systems (Briscoe et al., 1990).
CVM can be used to elicit values across the spectrum of
total economic value. It is generally regarded as the only
method for arriving at option prices and existence values.
Since there are few surrogate or implicit markets for these
values, indirect techniques relying on revealed preferences
are often of little use. One commonly cited exception is the
use of contributions to conservation organisations as a
surrogate market for existence or option values. There are,
however, a number of difficulties with this assertion, not
the least being getting at the actual reason people make
contributions. Option and existence values are discussed in
greater detail later on in this chapter.
Hedonic pricing
The hedonic pricing technique relies on the observation that
the value of non-marketed environmental services are
frequently incorporated into the prices of other marketed
goods and services. By disaggregating such market values
an economist may uncover the relative contributions of
valued attributes to human welfare. Although soil fertility,
scenic beauty or air- quality are not directly exchanged in
markets, hedonic pricing techniques enable economists
explicitly to value these services that are implicit in the
price of land and property, and wages.
The technique involves two stages. The first is relatively
straightforward, involving an econometric estimation of the
value contributed by the chosen environmental attribute to,
for example, property values. The second stage involves
working from this hedonic price equation back to the actual
demand curve. This second stage entails overcoming both
theoretical and practical obstacles. Basically, the output of
the first stage identifies the price for environmental quality
paid in a competitive market - not what the buyers are
willing to pay. In order to identify the WTP, analysts must
make simplifying assumptions about consumer preferences
or gather additional data on consumer preferences.
Applications of this complex and often ponderous technique
are scarce outside developed countries. The data
requirements are one very large drawback to undertaking
such studies in countries with a poor statistical base.
Travel cost method
The travel cost method is frequently applied to valuation
problems involving ecotourism and recreational services
derived from the environment. The technique requires
information on the expenditures by site visitors.
Aggregating the number of visitors by what it costs them to
travel to and from the site provides a surrogate market
indicating what people are willing to pay for access to the
site. Essentially, travel costs form a variable admission
price to the site. Those visitors from far away exhibit a
large WTP, while those from surrounding areas reveal a
low WTP. This relationship between distance and travel
costs can be used to estimate the benefits that visitors gain
by visiting the site. Drawbacks and potential obstacles to
the techniques involve unobserved travel costs, the question
of whether leisure time and travel to the site are necessarily
a cost and the fact that trips are usually multi-purpose.
Applications of the travel cost methods to protected areas
and other tourist and recreational sites in developing
countries are rare but likely to increase as the technique is
not overly demanding in terms of data or calculations.
In a recent application of the travel cost method Tobias and
Mendelsohn (1991) examined the willingness of local
tourists to pay to visit the Monteverde Cloud Forest
Reserve in Costa Rica. The research involved gathering
data from the reserve’s headquarters on the frequency of
ecotourist visits from each of Costa Rica’s 81 cantons.
Average distances from the major towns in each canton
were measured and then multiplied by an estimate of the
per kilometre travel costs in Costa Rica. Population density
and illiteracy were included alongside this travel cost as
variables that might explain the visitation rates for each
canton.
These results were used to calculate the ecotourism value
generated by the reserve. On average each visitor valued
the experience at US$35. The present value of such trips,
assuming constant flow of visitors and a real interest of 4%,
came to around $2.5 million. Because only one out of five
visitors to the reserve are Costa Ricans the total ecotourism
value is actually much larger. Foreign visitors are likely to
have far greater travel costs than local citizens, but are less
likely to be travelling to Costa Rica just to see the
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Biodiversity and Economics
Monteverde Cloud Forest. The authors confront this
difficulty by making the safe assumption that foreigners
value the experience as much as locals - leading to a present
value of $12.5 million. Since the total area of the reserve is
10,000ha the value per hectare for the reserve land is
estimated to be $1,250. When compared to the going price
for land surrounding the reserve of $30-$100 per hectare,
Tobias and Mendelsohn assert that expansion of the reserve
is called for on economic grounds. Their case would be
strengthened if the other direct, indirect, option and
existence values were included in the calculation.
Other techniques
A number of methods for deriving the value of
environmental resources exist. Although relatively inexact
these methods are often the second and third best techniques
that are actually used when time and money do not allow
for detailed research. Estimating how much would need to
be spent in order to prevent expected damage to
environmental quality is one way of valuing resource
degradation. Another way is to estimate the costs of
replacing the environmental asset that is degraded either by
the use of natural or man-made goods or services. This can
be accomplished by pricing available substitutes or the cost
of developing substitutes. The price of an environmental
improvement or cost of degradation may also be generated
by assessing the opportunity costs of the relevant action. A
final method of assessing the value of resources/resource
damage is to examine the cost of relocating economic
activity should the resource flow be disrupted.
LOSS OF BIODIVERSITY AS AN ECONOMIC
PROCESS
It is predictable that the application of economic processes
to natural resources will lead to substantial losses of
biological diversity. This is part of the general process
whereby humans continue to modify their natural
environment so that it can better satisfy their needs. To a
large extent, loss of biodiversity is a concomitant of this
value-maximising process.
However, it is also predictable that this process is unlikely
to target an adequate amount of diversity, because persons
deciding to convert their local environments do not consider
the global costs of so doing. From the economic
perspective, the biodiversity problem requires the regulation
of local development processes for the advancement of
global interests.
Conversions and loss of diversity
The economic theory of natural resources predicts that
much of existing diversity will be depleted. This is because
economics views the natural form of the resource as being
necessarily competitive with other forms in which humans
might hold these same resources: natural resources are
‘natural capital’ in contrast to ‘man-made capital’. Humans
make the choice of whether to hold the resources in their
original form, or to convert them to a modified form
(Solow, 1974).
2. Uses and Values of Biodiversity
From this perspective, human history has been one
continuing process of the conversion of natural resources
into more productive resource forms. For example, iron ore
is more productive in the form of a machine than it is as a
vein of sub-surface minerals. Therefore, the natural form of
the capital is altered to make it more useful, resulting in the
man-made form of the capital.
Similarly, a given hectare of land originally growing
diverse native grasses will be converted to another plant
form, such as wheat, because of the enhanced productivity
of this resource. That is, there is a choice to be made not
just between natural and man-made resources but also
between more and less productive forms of natural
resources. The biosphere can capture a limited amount of
useful solar energy, and it is now unavoidably a human
choice to determine which species will be used to perform
this task over much of the earth’s surface. Economics
indicates that humans will choose to channel this energy
only through those species which are most productive,
eliminating the others through this competitive process.
These conversions have been driven by two important
economic characteristics of resources: the relative growth
rates of what humans wish to consume and the relative
harvest costs. In economic terms, it is predictable that those
species which exhibit relatively high growth rates and low
harvest costs will displace many of the others.
Specialisation and loss of diversity
There are good reasons to believe that prevailing methods
of production are biased against the maintenance of a wide
range of diversity. This is attributable to the economies of
scale implied by the law of economic specialisation.
Biodiversity losses will not only result from the substitution
of the more productive resources for the less; these losses
will also occur by reason of the inertia that will develop
around the more productive forms of natural resources.
The law of specialisation is one of the first laws of
economics, developed by Adam Smith in the 18th century.
He observed that there tended to be increasing productivity
with increasing homogeneity in production methods and
processes. It is far less costly, in general, to create
thousands of units of an identical product than it is to make
smaller numbers of differentiated products. This is why
‘handcrafted’ goods are more expensive than factory
produced ones.
The cost differential is attributable to the application of
capital goods in the process of mass production. Once
capital is applied to the production of a particular good, it
usually becomes much less costly to produce. However, the
use of capital also implies homogeneity in the product. It is
the very essence of capital that it must be fine-tuned to the
production of goods of very specific characteristics.
In terms of biological products, the capital goods applied in
mass production are the chemicals and machinery of
intensive agriculture. These capital goods do not enhance
the general productivity of the biosphere; rather, they
increase its productivity by means of specialised substitution
of natural resources. The diverse resources of nature are
removed in favour of the specific resources for which
410
capital goods have been developed. Cultivators and
harvesters are developed to work in fields that are planted
with a single crop. Chemicals are fine-tuned to eliminate all
competitors of that crop.
The result of such specialisation is that an increasingly
narrow spectrum of species meets all of the needs of
humankind. A very small proportion of the thousands of
plant species which are deemed edible produce the vast
majority of the world’s food. The four major carbohydrate
crops (wheat, maize, rice and potatoes) feed more people
than the next 26 crops combined (Witt, 1985). This also
applies to animal protein sources. The tables of the Food
and Agriculture Organization list only a handful of
domesticated animals (sheep, goats, cattle, pigs etc.) which
supply nearly all of the non-fishery animal protein for the
vast majority of humans.
This concentration on a few useful species is occurring not
only because these are relatively productive and
manageable, but also because of the inertia resulting from
specialisation. The economies realised from mass
production continue to become greater as larger capital
goods (larger machinery, larger farms, more chemicals) are
employed, but this also implies increasingly homogeneous
production. This means that the gains derived from
specialisation also entail losses of diversity.
Globalisation and loss of diversity
The production of homogeneous capital goods also results
in increasing economies. Producing a wide range of tractors
and harvesters each tailored to a different crop is
inefficient. Making a single style of machine to be
employed the world over is the least costly method of
producing capital. The same applies to chemicals. It will be
less expensive to continue to fine-tune these to a few crop
species, and mass-produce these crops, than it will be to
produce chemicals adjusted to a range of different species.
Therefore, diversity losses do not occur only because of
relative differences in natural productivity, and because of
the inertia that develops around a given species once capital
goods are applied to its production, but also because of
inertia which develops around a particular type of capital
good. Once a particular species has been chosen for capital-
intensive production, it represents a commitment to a
particular technology and mode of production. As capital
spending becomes ever larger in regard to agricultural
production (e.g. biotechnology investments), it becomes
more important to increase the amounts of the specialised
species produced in order to be able to spread the fixed
costs of the investment, and to do this across both space
and time. Thus, the spread of intensive agriculture across
the world (including the ‘Green Revolution’) is predictable
as a method of spreading these fixed costs across space.
Specialisation and globalisation have gone hand-in-hand to
generate worldwide losses of diversity in the furtherance of
agricultural productivity.
Overshooting optimal diversity
In conclusion, much of the global loss of biological
diversity derives from the relative advantages of particular
species and particular methods of production.
From an economic perspective, the loss of some diversity
is inevitable. The issue at stake is whether the decrease
taking place at present is optimal in maximising benefit to
mankind, or whether it is in fact excessive (in economic
terms, whether the process will ‘overshoot’, or has already
done so). There are several reasons why loss of diversity
will tend to be excessive.
First, there is the possibility that current diversity is being
too readily traded-off for immediate gains in productivity.
At some point in time humans might decide that they would
prefer to have more than four sources of carbohydrate or
more than a dozen sources of protein, but by then diversity
might have been reduced to such an extent that it is no
longer possible. Similarly, it might be desired, if life-
sustaining income levels are achieved world wide, that the
world should contain more diversity to experience and
enjoy; this will not be possible if diversity has been over-
exploited.
Second, there is an increasing level of risk attached to a
strategy of specialisation. Diversity supplies insurance
against unforeseen events which specialisation does not
provide. There can be an increase in the average
productivity from conversions, but its variability might
simultaneously increase, and because of the increasing risk
of further conversions, the cost of each is not the same.
There is a cost involved in converting diversity that is felt
globally but not considered locally.
Third, the earth’s natural ecosystems are being altered at an
unprecedented rate, and at a rate far faster than our
understanding of them is advancing. Our knowledge of the
interactions between different parts of the biosphere is
particularly inadequate.
In summary, it is possible that some aspects of resource
conversion, while locally desirable, will have effects which
are undesirable from a wider perspective. This asymmetry
results from the element of ‘globalised value’ that attaches
to the remaining biodiversity. From the economic
perspective, the problem to be solved is how to bring this
‘external value’ into the equation when land-use decisions
are being made. To do so, it is necessary to acquire some
understanding of the nature of these values.
CURRENT USES OF DIVERSE RESOURCES
Introduction
The range of uses to which biological resources are
currently put was surveyed briefly in Chapters 25 and 26.
The intention here is to demonstrate how monetary value
can be attached to wild resources, by reference to two kinds
of use or consumption.
Diverse resources make substantial contributions to current
consumption in both a relative and an absolute sense. That
is, a notable portion of the world’s population relies for a
Significant part of its sustenance upon wildlife resources,
particularly in many developing countries (Prescott-Allen
and Prescott-Allen, 1982). In addition, there is significant
consumption of wildlife resources in those countries where
the vast majority of consumption does derive from
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Biodiversity and Economics
monocultures. Although the percentage value of these
resources in terms of the overall economy of the countries
is small, the absolute amount of value attached is very
large.
Even though wild resources are being replaced by
monocultures in the economic process of global conversion,
they are far from valueless, even in terms of their current
known use. It is important to recognise that it is the value
of wild resources relative to specialised resources that will
determine the extent to which wildlands will be converted.
If there is no added value from converting land from a
natural state, then the process of conversion will cease. One
of the important problems of biodiversity conservation
arises from the irreversibility of the conversion process.
This means that decisions by current generations regarding
the loss of biodiversity cannot in general be undone. Future
generations must live with these decisions even if their
values are different.
There is good reason to expect that the values of certain
diverse resources will systematically increase over time. It
has been argued that society’s preferences shift toward
natural resources and wilderness experiences, as wealth
increases and natural resources become scarcer. This is
evident in increased international tourism to places of
natural beauty, the developmentof ‘ecotourism’ and wildlife
encounters, and the willingness to pay for preservation.
Some societies have long held a preference for natural
products over synthetic varieties. For example, the Japanese
are renowned for their dissatisfaction with man-made
substitutes for wildlife products (Barbier et al., 1990).
These uses of wildlife resources are ways of expressing
preferences for the natural form of the habitat (and its
products) over the domesticated form. They are important
as means of countering the trend toward increased
conversion. There are problems to be solved with regard to
the use of wildlife resources, as there are with many kinds
of resources, but it is important to conserve more wild
resources now so that there is not a deficit of variety to
meet the needs and desires of future generations.
COMMUNITY USE OF WILDLIFE RESOURCES
State provision for wildlife protection can create conflicts of
interest between local communities and the protecting
bodies. The establishment of protected areas for wildlife is
often in direct conflict with the economic interests of local
communities. Many are denied access to resources that they
have traditionally exploited. Rural communities have not
only lost their traditional management and use rights to
local wildlife resources, but they may also bear the full
costs of crop damage because of migrating wildlife. The
situation has deteriorated in recent years, with rising rural
populations and increased poverty. Illegal encroachment,
hunting and harvesting are often the only available means
of securing subsistence and income. Where local
populations are not directly involved in these activities,
their alienation from wildlife resources means they have
little will to oppose the exploitation of these resources by
others. "The breakdown of traditional common-property
management regimes into virtual open-access exploitation
leaves rural communities with little means to enforce
sustainable management" (Barbier, 1990).
2. Uses and Values of Biodiversity
Table 27.1
OPTION 5 years
Game viewing with no 34.7
consumptive uses
Game viewing with 91.2
elephant cropping
Source: J. Barnes, Department of Wildlife and National Parks, Botswana
Note: * Values in million pula, discount rate 6%
Several programmes have attempted to counteract this by
diverting some of the revenue generated by sustainable
management of wildlife populations into the hands of the
communities in which these populations are concentrated.
Community based utilisation of African Elephants
Elephant culling has been undertaken in Zimbabwe since
1965, with the objective of controlling elephant numbers.
Revenues from the sale of ivory (legal and confiscated
illegal ivory), skins and meat are a natural byproduct of
such population control policies. These revenues fund the
Management programmes and compensate the local
communities for elephant damage. They also support anti-
poaching activities which protect the rents available from
the sale of ivory.
Revenue from tourism and culling (the sale of ivory, hide
and meat) generates a very persuasive argument in favour
of elephant conservation. The combined value of elephants
from the sale of meat products and tourism in Tanzania has
been estimated in excess of US$80 million. Elephants
constitute a major tourist attraction generating tourist
revenues in excess of $25 million annually; were
populations to recover sufficiently, sale of products could
yield an additional $10 million; illegal meat hunting
currently generates around $40-50 million annually. Not all
the values derived from each of these activities can be
realised simultaneously: sustainably managed populations
cannot be subject to illegal poaching (ITC, 1989).
Comparable figures for the projected values from elephant
utilisation in Botswana are presented in Table 27.1.
Although game viewing alone results in significant returns,
these are more than doubled if elephant culling is included.
The Nyaminyami Wildlife
Zimbabwe
Management Trust,
The Nyaminyami Wildlife Management Trust (NWMT) was
formed by the Nyaminyami District Council in Zimbabwe.
Its objective was to administer the management of wildlife
resources for the benefit of the local inhabitants. The Trust
established hunting and culling quotas for wildlife, pursued
anti-poaching measures, and set up two Impala Aepyceros
melamprus sites where herds could be sustainably managed.
It also licensed two safari operators and metered
compensation for economic losses incurred by residents as
the direct result of conservation practices.
Projected values from management of elephant resources in Botswana
NET PROJECTED VALUE *
10 years 15 years
98.1 160.6
198.4 288.9
Table 27.2 Nyaminyami Wildlife
Management Trust
revenue*
Revenue NWMT
Buffalo Range Safaris 148,349
Astra Wildlife 117,790
Mashonaland Hunters 6,048
Sub-total Safari Hunting 272,187
Cropping 1 11,554
Cropping 2 24,356
Sub-total Cropping 35,910
Meat and skins,
Kapenta fishing 11,256
Total Revenue 319,353
Recurrent expenditure
Cropping costs 1 -10,244
Cropping costs 2 -18,604
Wages and salaries -16,378
Transport and equipment hire -636
Vehicle maintenance/fuel/repair/insurance -5,829
Wildlife compensation -26,681 .
Kapenta licences -2,400
Advertising, publications, printing -4,341
Miscellaneous, bank charges -1,469
Total Recurrent Expenditure -86,582
Net Revenue 232,771
ZimTrust 20,093
Adjusted Net Revenue 252,864
Source: Adapted from Jansen, D.J. 1990. Sustainable Wildlife
Utilisation in the Zambezi Valley of Zimbabwe: economic, ecological
and political tradeoffs. Project Paper No. 10, WWF Multispecies
Project, Harare.
Note: * In Zimbabwe $; in 1989 Z$2.1 = US $1.
In 1989, the Trust earned Z$319,353 in wildlife revenues;
approximately 85% of this came from concession and
trophy fees paid by safari hunters, and the remainder from
sales of meat, skin and hides. Through additional
contributions from the ZimTrust, Z$20,093 to finance
recurrent expenditures and Z$191,683 for capital
expenditure, NWMT ran a surplus of Z$252,865 (Table
27.2). This surplus was distributed between a reserve fund
for capital expenditures (12%), levies retained by the
district council (10%) and the remaining 78% to be
channelled back into the communities, funding housing
projects, clinics, teaching, and recreational facilities. A total
of Z$198,000 was disbursed throughout the communities,
constituting about 15-20% of annual household incomes. In
addition, direct compensation amounting to Z$27,681 was
paid for crop and animal damage, offsetting the costs
associated with wildlife conservation. Meat and skins from
cropping were also sold locally at a subsidised price
representing an additional net gain to the local inhabitants
from enlightened wildlife management.
For programmes such as this to work effectively, the
revenues must be channelled back to the community. In
Zimbabwe, trophy hunting fees paid by operators to the
central government should have been redirected back to the
community through investment in schools and clinics.
However, only 57% of the nearly Z$6 million earned from
wildlife over the period 1980-1987 had been returned by the
end of 1987. The Zambezi valley project generated wildlife
revenues of Z$2.1 million between 1981 and 1986, but by
1987 only 44% had been returned to the districts.
Community use of the Vicufia, Argentina
The Vicuiia (Vicugna vicugna) is a wild camelid inhabiting
the puna, a treeless pastoral zone in the central Andes of
western South America. Vicufa have fine wool and are a
sought-after meat delicacy. They have been hunted for
centuries and the Incas are recorded as following sound
management practices in harvesting them.
In 1987, Vicuiia populations in the Laguna Blanca Reserve
(Catamarca province) were examined to assess their
potential contribution to the indigenous peasant economy.
This is primarily a subsistence economy, with a small but
increasing involvement in the market economy. The two
main sources of income are from sheep and llama spun
wool. The potential harvest of the Vicufia population was
estimated using simulation techniques, calculating the
maximum sustainable yield and the carrying capacity of the
area (Rabinovich et al., 1991). If the Vicufia population
were allowed to grow from its current size of 5,000 animals
to around 8,000, 15.2% of that population could be
harvested each year. The monetary value of each Vicuia is
estimated at US$64: $19 for the wool, $10 for the meat
(assuming a 20kg animal fetches $0.50 per kg) and $35 for
the hide. The estimated total income that could be derived
from sustainable management of the Vicufia is US$94,464
per year. This would provide an annual household income
to the peasant community of the Laguna Blanca Reserve of
almost US$1,000 if equally distributed among the 95
families.
ECOTOURISM
Ecotourism, or nature tourism, is just one component of the
tourism industry. A precise definition of tourism is elusive
because of its complex nature, involving a combination of
attractions, transport, accommodation, supporting facilities
and infrastructure. It is generally defined by its spatial
dimension (Pearce, 1989), and is thus often characterised by
criteria such as a minimum distance of travel or travel
involving at least a one-night stay away from home.
Tables 27.3 and 27.4 present data and projections on
worldwide tourist arrivals and receipts from the World
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Biodiversity and Economics
Tourism Organization (WTO). Note that these figures
pertain to cross-border tourism; according to WTO purely
domestic tourism may be worth ten times as much as the
$250 billion generated by international tourism in 1990.
While political and military events have strongly influenced
recent tourist movement the upwards trend in arrivals and
receipts during the past five years is clear. Over the 1985-
1990 period tourist arrivals grew by 6% and receipts by
over 16%. WTO forecasts through to 2000 envisage
continued growth but at a reduced rate of 4% for arrivals
and 8% for receipts.
Although Europe accounts for the bulk of the world’s
tourist arrivals (over 60%), Europe’s share of the receipts
is not as large, presumably because of short intra-European
stays. The fastest growing segment of the market is in Asia.
WTO predicts that Asian tourist receipts and arrivals will
exceed that of the growing American market by the year
2000. Meanwhile, the African market for tourism remains
a small fraction of the world market, accounting for just
three per cent of world arrivals and two per cent of world
receipts in 1990.
Table 27.5 presents data on arrivals in all regions from the
Americas and Europe. It reveals that almost 90% of
travellers from Europe go on holidays to Europe or North
America. Similarly, a majority of travellers from the
Americas either stay at home or go to Europe. Clearly
some of the North-North travel is ecotourism (in 1986 US
parks brought in foreign exchange worth $3.2 billion);
however, destinations such as East Africa, Central America,
and Southeast Asia, which are renowned for their wildlife,
are clearly of only marginal significance in the overall
tourism picture. This contrasts with the fact that the tropics
are very rich in biodiversity and the temperate latitudes,
including Europe and North America, much less rich.
Nonetheless, tourism revenues may be of great economic
significance to local economies, particularly in developing
countries that are popular destinations for nature tourism.
In this regard, Swanson (1991) has calcuiated that tourism
provides 9-13 % of exports from developing nations in sub-
Saharan Africa, South and East Asia and Latin America.
Lindberg (1991) reports that tourism in Kenya generated
$400 million; in recent years it has been Kenya’s largest
earner of foreign exchange. Dixon and Sherman (1990) put
tourism’s share of the economy in Caribbean nations at 15-
30%. Tourism can obviously provide a boost to local
economies, but how much of this revenue comes from
ecotourism?
Defining the exact meaning of ‘ecotourism’ is no easier then
agreeing on the coverage of the term ‘tourism.’ Lindberg
(1991) characterises nature tourism as being distinctly
different from large-scale, highly developed, ‘mass’
tourism. Sites that attract Lindberg’s ecotourist feature
natural attractions and a certain degree of solitude. Lindberg
estimated that of the $55 billion in tourism revenues
accruing to developing countries in 1988, ‘nature tourism’
brought in 4-22% of these revenues. Despite its relatively
small share of the market, ecotourism, like other ‘special
interest’ sections of the market such as cultural tourism and
adventure travel, is expected to outpace the general growth
of ‘mass’ tourism in the next decade (Dixon and Sherman,
1990).
2. Uses and Values of Biodiversity
Table 27.3 International tourist arrivals
1985 1986 1987 1988 1989 1990 1995 2000
WORLD 322,723 330,527 356,787 381,946 414,223 429,250 515,000 637,000
EUROPE 214,263 215,396 230,752 239,347 266,946 275,500 294,000 338,000
AMERICAS 58,728 62,894 67,986 74,991 78,456 84,000 103,000 128,000
AFRICA 9,805 9,488 9,986 12,646 13,604 14,000 23,000 32,000
ASIA 39,927 42,749 48,063 54,962 55,217 55,750 95,000 140,000
east 29,408 33,128 38,372 44,703 44,387 46,500
mid—east 7,979 6,890 6,984 7,379 7,775 6,000
south 2,540 2,731 2,707 2,880 3,055 3,250
Source: from World Tourism Organisation
Note: figures are given in thousands
Table 27.4 International receipts from tourism
1985 1986 1987 1988 1989 1990 1995 2000
WORLD 116,158 139,234 170,456 196,521 209,416 249,300 343,000 527,000
EUROPE 61,181 77,024 96,341 106,746 109,007 136,300 152,000 206,000
AMERICAS 33,314 37,383 41,982 49,632 56,600 65,900 95,000 146,000
AFRICA 2,601 2,993 3,687 4,625 4,479 5,000 10,000 14,000
ASIA 19,062 21,834 28,446 35,518 39,330 42,100 86,000 161,000
east 12,851 16,118 21,281 28,394 32,405 36,500
mid—east 4,811 4,036 5,311 5,233 4,944 3,500
south 1,400 1,680 1,854 1,891 1,981 2,100
Source: from World Tourism Organisation
Note: figures given in $US millions
Table 27.5 Tourist arrivals, 1988
REGION OF DESTINATION REGION OF ORIGIN
EUROPE AMERICAS
ARRIVALS % OF TOTAL ARRIVALS % OF TOTAL
EUROPE 206,482,068 88.61 17,627,702 22.25
ASIA 4.69 5.74
Eastern Asia 2,160,959 0.93 2,691,257 3.40
Southeast Asia 2,703,173 1.16 1,018,680 1.29
Southern Asa 1,059,245 0.45 254,362 0.32
Western Asia 5,007,702 2.15 586,395 0.74
AMERICAS 412 70.33
Northern America 7,300,070 3.13 36,988,699 46.69
Southern America 1,054,539 0.45 5,756,668 7.27
Caribbean 1,010,705 0.43 6,601,596 8.33
Central America 231 886 0.10 6,370,939 8.04
AFRICA 2.26 0.63
Eastern Africa 624,460 0.27 117,650 0.15
Middle Africa 38,904 0.02 5,802 0.01
North Africa 3,933,058 1.69 263,862 0.33
South Africa 259,970 0.11 61,787 0.08
Western Africa 416,808 0.18 47,750 0.06
OCEANIA 0.31 1.05
Aust. & N. Zealand 662,693 0.28 594,285 0.75
Melanesia 25,359 0.01 65,684 0.08
Micronesia 1,397 0.00 80,041 0.10
Polynesia 43,895 0.02 90,862 0.11
TOTAL 233,016,891 79,224,021
Source: from World Tourism Organisation
414
More precise estimation of the size of the ecotourism
market is a near impossible task which requires a far more
discriminating statistical base than is currently available in
the national-level figures given to the WTO. While some
activities can clearly be classified as ecotourism (eg. safaris
to view the Mountain Gorillas in the Pare National des
Volcans in Rwanda), much tourism defies such
disaggregation. Most tourism is motivated by a mixture of
cultural, historical, biological, geological and personal
attractions. Evidence of this is demonstrated by Table 27.6
which presents the results of a survey examining the
motives of tourists travelling to five countries in Latin
America. An additional problem in specifying the value of
ecotourism is determining which receipts should be
allocated to which type of tourism. The bulk of the receipts
for tourist expenditures do not occur at tourist sites such as
parks, museums and cultural festivals, but at hotels,
restaurants and for travel costs.
For these reasons, attempts to demonstrate the value of
ecotourism often focus on illustrating the importance of
charismatic species to the conservation of particular natural
sites. For example, the Parc National des Volcans in
Rwanda receives approximately US$1 million a year in
entrance fees, with an additional $9 million in indirect
benefits to the local economy (Lindberg, 1991). In this
case, the park’s survival depends entirely on one species:
the gorilla. If ecotourists were not enthralled by, and
willing to pay for, the prospect of a face-to-face encounter
with gorillas, the park’s natural habitat would doubtless
long since have been converted to other uses.
In other cases, tourists may be attracted by a range of
species. A number of studies have put rough figures on the
value of elephants and lions in Kenya. Western and Henry
(1979) found that each lion in Amboseli Park in Kenya
generated US$27,000 in tourism revenues, while elephant
herds in the same park produced $610,000 per year. In a
further study of the tourism value of lions in Amboseli,
Thresher (1981) calculated that over a 15-year period a
single lion would draw in $515,000 in foreign exchange.
More recently, Brown and Henry (1989) used contingent
valuation and travel cost methods to calculate that the value
of viewing elephants in Kenya is $25 million per year. Such
figures lend credence to the claim that the ecotourism value
of such species is far greater than their trophy value.
Barnes et al. (1992) point out that management of elephants
in Kenya should consider not just their ecotourism value but
their total economic value. The role of uncertainty in
valuing ecotourist use, the potential for large existence
values and the indirect value of the elephant as a species
with a large ecological role to play must also be
incorporated into an estimate of their total value. Of course,
conservation efforts in Kenyan national parks must also
consider the total economic value of lions, zebras, acacias,
and other resources. In order to make decisions that
maximise the net benefits to society, the total economic
value of the site and all its associated values (use and non-
use) must be considered. For example, although
conservation of gorillas in the Parc National des Volcans is
encouraged by the incentives of ecotourism revenues, there
are additional ecological and existence values that accrue to
415
Biodiversity and Economics
locals and the global community simply because the gorillas
and their habitat are protected.
A final, cautionary note must be added when discussing the
value of ecotourism and the receipts generated by the
tourism industry in developing countries. Tourism may
indeed have a macroeconomic multiplier effect (the indirect
and positive feedback effects brought on by the direct
expenditure of tourist monies); however, the extent of such
a multiplier will depend on the funds remaining in the local
economy. If expenditures on tourist hotels and restaurants
are promptly spent on imports or repatriated by foreign
companies, there will be no ‘multiplying’ effect. Pearce
(1989) reports that small Caribbean and Pacific nations may
lose half of their gross foreign exchange earnings to
expenditures on tourism-related imports. The World Bank
has estimated that developing countries lose 55% of gross
tourism revenues in such leakage (Boo, 1990). The lower
the availability of locally produced goods and services used
by tourists the worse this leakage becomes. Thus,
ecotourism is not a panacea that guarantees wise and
effective use of biological resources. If the revenues of
ecotourism do not accrue to national park systems or local
communities, there will be little economic incentive for
investment in the recurring costs of conservation activities.
Table 27.6 Reasons for selecting
travel destinations in Latin
America
REASON RESPONDENTS %
Natural History 167 38.3
Sightseeing 161 36.9
Visiting friends and/or relatives 132 30.3
Sun, beaches, entertainment 130 29.8
Cultural/native history 102 23.4
Business/convention 87 20.0
Archaeology 63 14.4
Source: Boo, E. 1990. Ecotourism: the potentials and pitfalls. World
Wildlife Fund, Washington.
Note: Total number surveyed = 436
EXISTENCE VALUES
Existence values are those benefits that are completely
disassociated from the use of a resource. Existence benefits
occur when people are willing to pay simply for the
pleasure they derive from knowing that particular species or
habitats continue to exist, irrespective of any plans they
may have to hunt, observe or otherwise use these biological
resources.
A range of terminology is used in referring to existence
values, including bequest, stewardship, vicarious and
intrinsic values. If the motivation behind these expressions
of value is to preserve the resource for future use, then the
criterion of non-use is not met and these values are better
considered as use values. An additional source of confusion
and debate is the status of so-called ‘intrinsic’ value.
Interpreted to mean the value accruing to species other than
Homo sapiens, this value is outside the scope of economic
analysis which is based solely on the expression of human
preferences.
2. Uses and Values of Biodiversity
Existence values may accrue to people in both the
developed and the developing world. Unfortunately, as
Table 27.7 demonstrates, the results of empirical research
to date comes mainly from the developed world, in
particular the USA. While casual observation may lead to
the expectation that existence values are a ‘luxury of the
rich’, further empirical work is needed to deny or confirm
this hypothesis. While many economists agree that people
are willing to pay for the mental satisfaction of knowing
species and habitats exist, the psychological nature of
existence values has so far defied the emergence of serious
theoretical or analytical approaches on these values. With
little in the way of theory to guide empirical investigations,
the insight gained from case studies of existence values is
often limited to a mere examination of their size relative to
use values or option prices.
A study of preservation bids (synonymous with option
prices) for Bighorn Sheep and Grizzly Bears in Wyoming
by Brookshire et al. (1983) provides an excellent
illustration. Using survey questionnaires (the contingent
valuation method) the authors measured the willingness of
prospective hunters to pay for hypothetical future permits to
hunt Bighorn Sheep and Grizzly Bears. In addition the
questionnaire also identified existence values and observer
preservation bids. The bids of respondents indicating that
they would neither hunt nor directly observe the animals
were taken to reflect existence values. The results of the
study revealed a range of hunting bids from just under $10
to almost $30. Observer option bids for the two species
were in the vicinity of $20. Existence bids for the Bighorn
Sheep were in the $7 range while those for Grizzly Bears
averaged $15. This study reveals that existence values for
species may be of the same order of magnitude as option
prices for such direct uses as hunting and game-watching.
Table 27.7 Empirical measures of
existence values
VALUE PER ADULT RESPONDENT
IN MID-1980s (US$)
Animal Species
Bald Eagle
Emerald Shiner
Grizzly Bear
Bighorn Sheep
Whooping Crane
Blue Whale
Bottlenose Dolphin
California Sea Otter
Northern Elephant Seal
Natural Amenities
Water quality (S Platte River Basin)
Visibility (Grand Canyon)
Additional park facilities (Australia)
Sources: Pearce, D.W. 1990. An Economic Approach to Saving the
Tropical Forests. LEEC Paper DP 90-06. IED, London. Majid, I.,
Sinden, J.A. and Randall, A. 1983. Benefit evaluation increments to
existing systems of public facilities. Land Economics 59:377-392.
In a survey of the willingness to pay for additional park
facilities in Australia, Majid et al. (1983) demonstrated that
the existence values for habitat are also of a comparable
size to their recreational use values. The initial survey
questions asked respondents how much they would pay for
recreational use benefits and total benefits generated by a
list of current and proposed facilities. As a measure of
existence value the authors calculated the difference
between the willingness to pay for recreational site visits
and the total willingness to pay for each site. The results for
all parks indicated that the total benefits were roughly twice
as big as the use values - thus existence values were judged
of equal value to recreation values.
Table 27.8 Gifts to surveyed environmental/wildlife organisations
TOTALS
Nature Conservancy
WWF and the Conservation Foundation
Ducks Unlimited, Inc.
Sierra Club
Natural Resources Defense Council
National Audubon Society
National Arbor Day Foundation
New York Zoological Society
Sierra Club Legal Defense Fund
World Resources Institute
American Farmland Trust
International Fund for Animal Welfare
Resources for the Future
Animal Protection Institute
American Humane Association
American Forestry Association
Clean Water Fund
Adirondack Council
American Rivers
Trout Unlimited
Earth Island Institute
Rainforest Alliance
Soil and Water Conservation Society
Farm Sanctuary, Inc.
Alliance for Environmental Education
Wildife Habitat Enhancement Council
Lake Michigan Federation
Animal Rights Network, Inc.
American Cave Conservation Association
Peace Garden Project
US$,000 US$,000 PERCENT
1989 1990 CHANGE
208,907 273,385 3
48,963 85,527 75
33,465 42,438 27
25,501 29,674 16
21,908 28,718 31
12,524 13,821 10
10,174 11,094 9
8,126 11,045 36
17,073 9,531 —44
5,973 6,833 14
5,240 6,336 21
2,716 5,195 91
3,912 4,555 16
2,651 2,948 1
435 2,607 499
1,992 1,903 -4
909 1,816 100
719 1,607 124
1,178 1,542 31
1,728 1,502 -13
1,180 1,309 11
1,007 1,026 2
254 798 214
388 390 1
165 346 110
19 186 879
152 182 20
133 165 24
200 118 -41
87 108 24
135 65 —-52
Source: AAFRC Trust for Philanthropy
416
But how are these existence and preservation bids actually
expressed in the real world? One way of expressing these
desires for the benefits of species and habitats is to make
donations to organisations that conserve biological resources
and biodiversity. Table 27.8 provides figures on such
philanthropic giving in the USA over the 1989-1990 period.
The data - as would be expected - reveal an upward trend
in overall giving to the organisations surveyed by the
AAFRC Trust for Philanthropy. Table 27.9 reveals figures
for the total charitable contributions to environmental and
wildlife causes in the USA in the context of total giving.
While $2.3 billion is a substantial sum of money, the
amount donated to the environment pales beside that
donated to other philanthropic causes. While environmental
giving has registered growth of 9%, 11% and, most
recently, 24% per year in real terms, the fact remains that
the average donation came to roughly $10 per person in the
USA in 1990.
Table 27.9 Charitable contributions in
the USA
Total Funds in 1990: $122.6 billion
Destination of Funds %
Churches and synagogues 53.7
Education 10.1
Human services 9.6
Arts and culture 6.4
Public benefit 4.0
Environment 1.9
International 1.8
Undesignated 4.4
Sources of Funds %
Individuals 83.0
Bequests 6.4
Foundations 5.8
Corporations 4.8
Source: AAFRC Trust for Philanthropy. 1990. Giving USA, NY.
A critical ingredient of CVM studies is the extent to which
tespondents are informed about the object of the studies
and, correspondingly, how much information is disclosed
during the survey process. Samples etal. (1986)
investigated the effects of information disclosure on
preservation bids for endangered species. Although
preservation bids may be interpreted to have both use and
non-use components, the authors assumed that the
endangered status of the species would cause responses to
reflect mainly on the value of ‘saving’ the species as there
was little real prospect for ‘using’ species close to
extinction.
THE VALUATION OF DIVERSE ECOSYSTEMS
Introduction
A particular cause for concern arising from the conversion
of biological systems is the problem of accumulated losses
of unknown ecosystem values.
Rational decisions about the conversion of one system to
another should involve an assessment of relative values;
417
Biodiversity and Economics
typically, however, the only values that are included in that
comparison are the appropriable ones. If a person or group
cannot capture that value, it is unlikely to be considered
important. However, many of the benefits of biological
systems flow not to any one particular individual or group
but to the community at large. Attributes of forests, such as
oxygen production and carbon fixation, are unlikely to stop
a logger from acting, even though these are very important
characteristics. The total economic value, including these
non-appropriable values, of biological systems must be
entered into the calculation if the optimal amount of
diversity is to be conserved. This is unlikely to occur at any
time in the near future, simply because we do not have the
capacity to do so. There is, therefore, good reason to
preserve some parts of the world’s diversity in general
recognition of the global public goods that it provides even
if these cannot be assigned a value.
THE VALUE OF TROPICAL FORESTS
The fact that tropical forests have value is not disputed:
they are a source of ecological benefit and material wealth
(see Chapter 20). In order to understand the consequences
of decisions made about different possible uses of the
forest, it is necessary to quantify and rank the values under
these different uses.
The method of system valuation
The purpose of valuation techniques is to correct those
prices that do not correspond to the ‘true’ economic values
and to calculate prices for those assets that are not valued
at all (Maler, 1989). The concept of total economic value
(TEV) offers a unified approach to the valuation of tropical
forests. This concept is based upon the idea that it is
possible in certain cases to disaggregate the flow of goods
and services from environmental resources, and then to
assign monetary values to these discrete functions.
Some of the component goods from forests are traded in
markets, and in these cases the market price provides an
indicator of social value. But this is so only if markets are
perfectly competitive and complete; then, the prices arrived
at will reveal the correct marginal valuations of those goods
and services exchanged. Otherwise, it is necessary to
compute a shadow price, i.e. a price that differs from the
market one but corresponds more closely to social value.
However, in many cases environmental goods cannot be
traded in the marketplace, hence their value is not directly
revealed. In this case other methods must be employed to
gauge their value and capture how that value alters with
different uses.
Therefore, the object of environmental valuation is the
performance of these three tasks:
the segregation of a unitary system into discrete
components
the valuation of those components that are not traded in
markets
the correction of market values, where these differ from
social values.
2. Uses and Values of Biodiversity
Table 27.10 The concept of total economic value* in
a tropical forest context
USE VALUE + NON-USE VALUE
DIRECT + INDIRECT + OPTION + {QUASI + EXISTENCE
VALUE VALUE VALUE OPTION VALUE
VALUE}
Sustainable
timber
Non-timber Nutrient Future direct Forests as objects
products cycling and indirect uses of intrinsic value,
as a bequest, as a
Recreation Watershed gift to others, as
protection a responsibility
(stewardship).
Medicine Air quality Includes cultural
Plant genetic Micro-climate
resources
Education
Human habitat
and heritage values.
Source: Pearce, D.W. 1990. An Economic Approach to Saving the Tropical Forests. LEEC Paper DP 90-06. ITED, London.
Notes: Direct Value refers to those benefits that can be observed being consumed, although their consumption might not yield a meaningful price
which can be assigned to that benefit. Indirect Value refers to those benefits that are not observed being consumed, but that are known to be
essential to the preservation and maintenance of ecosystems. Option Value is the value placed on securing the future consumption of goods and
services yielding direct and indirect value.
Quasi-Option Value is the value of learning about future benefits that would be precluded by
development or irreversible change of the forests today. This takes account of the fact that current valuations are circumscribed by current knowledge
of forest functions. Existence Value is that value placed on an environmental asset independent of its current or future ‘usage’. This incorporates
the innate value of the forest in situ.
Table 27.10 outlines the components of TEV with reference
to tropical forests. Trade-offs occur between the different
types of uses, direct and indirect: the supply of hardwood
might diminish the amount of protection offered to
watersheds by the root network of the trees; forest areas
devoted to recreational facilities might displace indigenous
peoples. We cannot simply add the components of TEV to
obtain a measure of the ecological wealth of the forests.
Direct use values in tropical forests
Timber
Logging can be consistent with forest conservation if the
forest lands are managed sustainably. The limitation to this
approach to timber valuation is that market prices generally
diverge from shadow prices (those that reflect the true
opportunity cost of the good or service). In particular, if the
forest is being logged without well-defined property rights
(without care for its future flow of timber), then it is likely
that the price of timber does not reflect the full amount of
resource rent that is available. The timber price might then
reflect only the social value of the labour and capital inputs
into its production, not the value of the timber itself. In this
case, the market price must be corrected to reflect the true
social value of the resource.
The management of all environmental resources is tied
closely to the problem of the valuation of those resources
over time. Given that one of the main reasons to conserve
today is to preserve the resource for tomorrow, it is
important to value a resource not at a single moment but
over a period of time. This allows the value of unused
418
* Total economic value = use value + non-use value
resources to come into the calculation. When future values
are combined with current values, it is important to take
into consideration the relative weights to ascribe to the time
periods. Usually, a discount is applied to future period
values, because of the uncertainties involved. The net
present value calculation is very sensitive to the discount
rate employed to convert the stream of future benefits into
a single value.
Discounting enables the economist to represent the value of
a resource or asset in terms of the flow of income deriving
from that asset over a specific time span. The discount rate
reflects the greater importance attached to current,
compared with future, consumption and the trade-off
between them.
One important study (Table 27.11) examined the
comparative present values of an Indonesian forest, given
a range of possible uses. It demonstrates the comparability
of returns available from a range of different forest
management practices for timber production. In addition, it
also illustrates the method of present valuation of the
production capability of a hectare of forest land.
The impact of utilisation on environmental quality depends
on how the forest lands are altered as a result. SAW and
PULP may threaten existing tropical forest-lands because
they are typically concentrated among uniform plantings of
non-indigenous species. However, where these plantations
occur on previously unforested land they may still
contribute to ‘carbon-fixing’ and have positive net
environmental worth.
Table 27.11 Profitability of logging at
different discounts
REGIME DISCOUNT RATE
5% 6% 10%
TPI 2,705 2,409 2,177
CHR 2,690 2,593 2,553
INTD 2,746 2,203
PULP 2,926 2,562
SAW20 2,419 2,278
SAW10 2,165 2,130
Source: D.W. Pearce. 1987. Forest policy in Indonesia. unpublished
memorandum. World Bank.
Notes: Net present value US$1,986/ha. TPI - selective cutting regime
in which only those trees over 50cm in diameter at breast height are
harvested. CHR - complete harvesting and regeneration; all
merchantable trees are harvested then the cleared land undergoes
natural or enriched regeneration. INTD - intensive dipterocarp
management, following a plantation approach on clear-felled land.
PULP - when plantations of fast-growing trees are harvested for wood
pulp. SAW - refers to saw timber plantations where the trees are cut
at 10/20 years respectively.
The table reveals that were an estimate of the net worth of
a project to be based solely on financial profitability it
would be preferable to encourage rapid-growth plantations
for pulp production. The more sustainable selective cutting
regime is only favoured at the lowest discount rate of 5%.
Therefore, although there is no clear-cut advantage to large-
scale alteration of the forest environment during utilisation,
the financial incentives from logging alone might encourage
this to happen. Focusing on this single use of the diverse
habitat can lead to the conclusion that habitat conversion is
economically optimal. This conclusion may not be the case,
even when only logging is being considered. In many
instances the conversion of forest lands is only financially
profitable when considered in combination with government
subsidies that encourage the same.
Non-timber products
This conclusion can be altered quite dramatically by the
introduction of a wider range of goods and services into the
analysis. Non-timber forest products are often a vital source
of foreign exchange earnings and revenue. They are also
essential to the rural household economy (de Beer and
McDermott, 1989). In many parts of Southeast Asia, the
tural population depends heavily on forest products for their
daily needs. It has been estimated that in Southeast Asia at
least 27 million individuals rely on the forests to satisfy
their nutritional, fodder, fuelwood, and _ shelter
requirements.
These products are seldom exchanged or sold, so care must
be taken in valuing their contribution to the rural economy.
It is possible for this purpose to use the value of the effort
expended as a surrogate. This can be done by calculating
the ‘cost’ of the labour inputs that are applied in gathering
and harvesting the non-timber products. In Thailand it was
estimated that the one million families who are forest-
dwellers devote about 180 person-days a year to collecting
forest-food, whilst the three million families located on the
periphery of forests spend about 60 person-days harvesting
food. This can be taken to indicate that these resources are
valued by forest-dwellers at an amount up to half a year’s
419
Biodiversity and Economics
salary (assuming that the hours spent gathering the
fuelwood could be otherwise redirected to the labour
market). In many cases the forest products represent
substitutes for goods that can be bought in local markets
and therefore the market values of these substitutes can be
aggregated to provide a monetary equivalent of the forest-
products. Of those products that are themselves marketed,
such as rattan, nuts, fruit etc., values may be more easily
discerned. Non-timber forest products also contribute
greatly to the national economy. They generate
employment, foreign exchange earnings, trading and
processing revenues, consumption and import substitution
opportunities.
In Thailand it was estimated that there were 200 rattan
furniture manufacturers operating small-scale cottage
industries which produce goods mainly for the domestic
market. In 1987 the US dollar value of exports totalled
US$29.1 million (Anon., 1988). Thailand also exports
finished bamboo products: handicraft export values vary
from US$212,413 (Ministry of Commerce) to about
US$3 million (a questionably large figure). Whilst attempts
to value bamboo exports produce varied results, bamboo is
nevertheless a widely-used product of great importance to
the national economy.
Ecotourism values
Tropical forests are also valued for their recreational
benefits. In Costa Rica, Ecuador, Philippines and Thailand
tourism is a vital source of foreign exchange earnings,
generating more revenue than the export of timber and
timber products.
The travel cost method is often employed to value the
benefits derived from tourism. Direct costs of access,
package tours, and hire of transport can all be regarded as
components of the overall value attributed to recreational
use. This approach has been used extensively in developed
countries to value the provision of recreational goods and
services. The methodology rests on the proposition that
observed behaviour can be used to derive a demand
function for non-marketed environmental goods and
services, regarding travel costs as surrogates for variable
admission or access costs. For example, European package
‘explorer’ holidays to the Peruvian Amazon cost about
$2,300 per person for 20 days.
Medicine and plant genetics
Tropical forests provide the habitat for a great variety of
species. The legal export of hides and skins, genetic
materials, spices, and oils provides many developing
economies with revenue. Where these products are traded
we may estimate the value of forest byproducts.
A very important product of forest diversity is its plant
varieties and the special information on chemical use that
these represent. Plant-based pharmaceuticals are a vital
source of foreign exchange earnings. In 1979 Thailand
exported medical plants and spices valued at US$17 million,
consuming about US$20 million domestically (Anon, 1981).
However, not all of the potential rents deriving from the
sale of these products are captured. In the case of
2. Uses and Values of Biodiversity
pharmaceuticals, calculation of the market value of drugs
bought does not yield an estimate of the full value of the
plant source because the value reflects not only the drug
manufacturers’ willingness to pay, but also the consumers’
net gains from the use of this plant. Here, the market price
is probably a poor indicator of the actual social value of the
good; it needs to be adjusted upwards.
Indirect use values
Whilst some forest conversion in the tropics provides
farmers and ranchers with valuable new tracts of land,
much leaves only degraded soils unsuitable for sustained
agricultural production. The loss of tree cover in watersheds
increases flooding, erosion, soil-leaching, and downstream
sedimentation. In semi-arid areas, deforestation depletes
essential organic matter, exposing the soils to wind and
water erosion. There is a very significant loss of ecosystem
function and the benefits which these systems render, both
on and off site.
The damage incurred as a result of the removal of forest
cover may provide an estimate of the value of watershed
protection. The loss of revenue because of declining soil
fertility, decreased freshwater fish yields as the result of
increased sedimentation, and reduced local rainfall can all
provide a measure of the indirect use values that accrue to
forest conservation.
Defensive expenditures designed to mitigate against the
effects of the loss of forest cover can also provide an
economic value for indirect usage. Such expenditures
include the cost of building levees, windbreaks, the
application of fertilizers, and increased irrigation
requirements. These, however, are undertaken with the
implicit assumption that the benefits from replacement
exceed the costs of deforestation. For if they did not, it
would not have been rational to deplete forest cover in the
first place.
Where forest cover is interrupted, nutrients are released into
the hydrological cycle. In general there is a net nutrient
outflow which can in itself pollute local river systems and
that greatly reduces the productive capacity of the cleared
land.
In growing, forests fix carbon dioxide through the process
of photosynthesis and give off oxygen. Once grown, there
is no net exchange of carbon and oxygen, mature forests are
described as being in carbon equilibrium, and in this state
they release as much CO, as they absorb. Deforestation
releases CO, (and other greenhouse gases such as methane)
into the atmosphere, contributing to the greenhouse effect.
In valuing the carbon-fixing properties of a tropical forest,
we must be careful not to double-count. Whilst preservation
ensures that the damage associated with carbon release is
averted, forest clearance results in a net debit. However, it
would be inappropriate to ascribe both a positive value to
carbon-fixing and a negative one to forest clearance in an
evaluation of the net benefits deriving from conservation.
The calculation is sensitive to the method of forest clearance
and the subsequent use to which the timber or forest lands
420
are put. If the forest is clear-felled and all the timber is
used to make durable wood products (housing timbers,
furniture etc.), then deforestation may cause little CO,
release because much of the carbon will remain contained
in the timber products. However, clearance through a
‘slash-and-burn’ approach will release all carbon contained
by the forest, with no offsetting gain from the productive
use of the forest timber.
Non-use benefits
Most attempts to develop existence values (those not related
to functional requirements) rely on the contingent valuation
approach, which reports the ‘willingness to pay’ of
individuals for environmental goods or services. To date
there have been no studies relating directly to the existence
value of tropical forests.
Cost-benefit analysis: the Korup Project, Cameroon
The following example illustrates the type of calculations
that might be undertaken to elicit a value for the net benefit
of a particular forest in situ. The aims of the Korup project
are to promote conservation of the rain forest in Korup
National Park in Southwest Province, Cameroon. It was
undertaken on behalf of the Government of Cameroon and
the World Wide Fund for Nature.
The project chose to evaluate the flow of benefits from
conservation options. The net benefits deriving from
sustained forest and subsistence use, tourism, genetic
materials, watershed protection, soil fertility maintenance,
and flood control are compared with the opportunity costs
of forestry and other development options (Table 27.12).
The opportunity costs measure that value of timber earnings
forgone by the preservation of the forest. The direct
benefits attempt to place a value on the sustained forest use
beyond the year 2020 when the forest would have
disappeared had it continued to be managed under the
current regime. They also give a figure for the replacement
of the subsistence production of the resettled villagers; the
value of tourism; the minimum expected genetic value of
the forest resources, etc. The induced benefits value the
contribution the project makes to agricultural productivity
and forest activities in the locality of the forest.
The final figure is then adjusted to reflect the net positive
contribution of the external funding to Cameroon, the fact
that Cameroon will be able to realise only 10% of the
genetic value through the operation of patents and licensing,
and that some of the watershed benefits accrue to Nigeria
and not to Cameroon.
Conclusion
The forest represents a wide range of values, from timber
to carbon-fixing. Incorrect decisions about use will always
be made if any one of these uses is considered in isolation
from the others.
Valuation becomes increasingly difficult as the use becomes
more removed from the marketplace. Thus, carbon-fixing
Table 27.12 Cost-benefit analysis: the
Korup Project
Direct costs of conservation -11,913
Opportunity costs
Lost stumpage value - 706
Lost forest use - 2,620
- 3,326
Direct benefits
Sustained forest use 3,291
Replaced subsistence production 977
Tourism 1,360
Genetic value 481
Watershed protection of fisheries 3,776
Control of flood risk 1,578
Soil fertility maintenance 532
: 11,995
Induced benefits
Agricultural productivity gain 905
Induced forestry 207
Induced cash crops 3,216
4,328
NET BENEFIT - PROJECT 1,084
Adjustments
External trade credit 7,246
Uncaptured genetic value - 433
Uncaptured watershed benefits - 351
NET BENEFIT - CAMEROON 7,545
Source: Ruitenbeek, H.J. 1989. Social cost-benefit analysis of the
Korup Project, Cameroon, prepared for the World Wide Fund for
Nature and the Republic of Cameroon, London.
Note: NPV £,000, 8% Discount rate.
values and nutrient cycles are real sources of value, but
very difficult to quantify. The Korup study demonstrates
that a careful attempt to derive these values indicates they
are very substantial indeed. The tropical forest resources
will be depleted if their entire range of values is not fully
Tecognised and integrated into decision-making by
individuals and governments.
THE VALUE OF WETLANDS
Wetlands are areas of land that remain waterlogged for a
substantial period of the year (see Chapter 22). Tropical
wetlands cover 2.64 million km? world wide whereas
wetlands in temperate and boreal regions occupy about 5.72
million km?. They support a wide variety of plant and
animal species restricted to such environments. Wetland
ecosystems are among the most threatened of all
environmental resources. Much of the physical loss of
wetland area has been because of the conversion to
industrial, agricultural and residential use. However,
qualitative degradation can occur in more subtle ways:
through discharge, effluent, and mechanical interference to
water flows. Wetlands are acutely vulnerable to damage
caused by activities located a considerable distance from the
wetland site but within its drainage basin.
As with tropical forests, the functions performed by wetland
systems are diverse. The structural components of wetland
systems (flora and fauna) are considered as stocks, whereas
Biodiversity and Economics
the ecological functions can be regarded as flows (services
that the wetlands yield over time).
An ecosystem is both a set of constituent characteristics and
the sum of these components. In many cases, the value of
the sum of the components is greater than the value of the
individual components alone. This is because some of the
functions of an ecosystem are able to continue only when
some significant proportion of the components are present.
Once some certain threshold is passed, the effect is to lose
these synergistic values. Therefore, the task of valuing an
ecosystem involves both the valuation of the components
and the identification of the synergism they generate.
A study of the Petexbatun wetlands in Guatemala provides
an indication of the range of values available at a single
wetland site. These values include direct use values, from
the generation of fisheries and wildlife habitat for example.
Less evidently, this wetland also provides a wide range of
indirect use values, by, for example, recharging inland
groundwater supplies and providing a buffer for flood
control. Finally, there are also the inappropriable values
represented by a wetland as a dynamic and diverse
biological system; although this is a non-use value, it is
probably one of the most important roles of the wetland.
The wide range of use and non-use values represented by
this single wetland are set out in Table 27.13.
Table 27.13 Wetland values:
Petexbatun, Guatemala
DIRECT INDIRECT NON-USE
Components
Forest resources eco
Wildlife resources e
Fisheries ee
Forage resources ee
Agricultural resources ee
Water supply eco
Functions
Groundwater recharge/
discharge e
Flood and flow control eco
Shoreline/
bank stabilisation eee
Sediment retention e00e
Nutrient retention e/ee
External support eco
Recreation/tourism e
Water transport eco
Attributes
Biological diversity ee eo eo
Uniqueness to culture/
heritage e
Source: Barbier, E.B. 1989. The Economic Value of Ecosystems: 1
tropical wetlands.
Notes: © = low e@ = medium eee = high
Case study: the Hadejia-Jama’are floodplain, Nigeria
Coherent policy determining the use and exploitation of
wetland resources requires that decision-makers have
available to them a set of shadow prices and values which
reflect the total economic value of these resources under
various management regimes.
2. Uses and Values of Biodiversity
One approach to valuing the wetlands is exemplified by the
case of the Hadejia-Jama’are floodplain in Nigeria. The
Hadejia-Jama’are wetlands lie in an area of confused
drainage between Hadejia (Kano State) and Nguru and
Gashua (Borno State), where the Hadejia and Jama’are
rivers flow across a fossil plain of late Quaternary sand
dunes. These wetlands provide essential income and
nutritional benefits for the regional inhabitants. They
constitute a source of fuelwood, fishing, grazing, and
agricultural opportunities. It is not only those located on the
periphery of the wetlands for whom this natural resource is
important. The floodplains provide dry-season grazing for
semi-nomadic pasturalists and agricultural surpluses for
Kano and Borno states, as well as educational and scientific
benefits. They also provide a natural habitat for migratory
and resident bird species.
However, the wetlands are shrinking as the result of
prolonged drought coupled with upstream water
developments which divert water flowing into the
floodplains. The Hadejia-Jama’are wetlands comprise dry
farmland and savanna, open reaches of water, swamp and
seasonally-flooded grassland. Agricultural practices vary
according to the terrain and comprise dryland agriculture on
the better drained sands together with various forms of
wetland cultivation, and seasonal grazing and fishing in the
more waterlogged soils and permanently flooded stretches.
The region experiences a single short wet season (May to
September); consequently the growing season for rain-fed
crops is short. Additionally, the region is characterised by
extreme rainfall variability, producing a high variance in
agricultural production. River flows are also highly
seasonal, with the timing, extent and duration of flooding
depending on the seasonal flood of the rivers and the height
of the water table beneath the plains. Thus the area and
nature of the wetlands also vary.
Direct use values
The direct uses of the floodplains encompass: fuelwood
collection, grazing of floodplain pastures, floodplain
agriculture and fishing, recreation, and transport.
The total cultivated area in the Hadejia-Jama’are floodplain
is estimated at approximately 230,00ha, of which roughly
77,500ha are cultivated in the dry season, and 152,500ha in
the wet season. The current annual net benefits from 14
agricultural crops grown in the Hadejia-Jama’are floodplain
have been estimated (Table 27.14).
Fishing is concentrated in approximately 100,000ha of
flooded land. Roughly 73,150 rural households in the
floodplain were estimated to fish throughout the year; 12%
of these households contained people whose main activity
was fishing, 21% were dry season fishing households, 15%
wet season fishing households and the remaining 52%
comprised households that only fished at fishing festivals.
Table 27.14 Agriculture: net benefits from the Hadejia-Jama’‘are floodplain, Nigeria,
1989-1990
CROP AVERAGE FINANCIAL ECONOMIC
OUTPUT PRICE PRICE
(Tonnes) (N/Kg) (N/Kg)
Total 7 281,955 26,710.50 19,092.72
Tradeable '
Rice 22,335 4,770 3,144.90
Wheat 43,350 4,010 1,382.88
Soyabeans 6,000 3,310 2,137.50
Non-Tradeable 2
Sorghum 50,315 720 612
Maize 15,705 842 715.70
Groundnuts 3,855 4,980 4,233
Millet 50,415 842 715.70
Cow-Peas 25,035 3,310 2,813.50
Tomatoes 15,955 662.50 563.13
Onions 1,1925 662.50 563.13
Peppers 32,400 1,336 1,135.60
Sweet Potato 2,925 600 510
Aubergine 1,740 662.50 563.13
Pumpkins 141,500 3 2.55
Net economic benefits per hectare: 239
(agricultural area 230,000ha)
FINANCIAL ECONOMIC NET ECONOMIC
BENEFITS BENEFITS BENEFITS
(N‘000) (N‘000) (N‘000)
563,104 366,455 54,968
106,538 70,241 10,536
173,834 59,948 8,992
19,860 12,825 1,924
36,227 30,793 4,619
13,224 11,240 1,686
19,198 16,318 2,448
42,449 36,082 5,412
82,866 70,436 10,565
10,570 8,985 1,348
7,900 6,715 1,007
43,286 36,793 5,519
1,755 1,492 224
1,153 980 147
4,244 3,607 541
Source: Barbier, E.B., Adams, W.M. and Kimmage, E. 1991. Economic Valuation of Wetland Benefits: the Hadejia-Jama’are floodplain, Nigeria.
Notes: Values in Naire per hectare; N7.5 = US$1 ' The economic prices of all tradeables are the c.i.f. import (border) prices converted at the
official 1989 exchange rate N7.5 = US$1.? Non-tradeables are defined as crops whose prices exceed f.0.b. export prices but are less than the c.i.f.
import prices. The economic prices of all non-tradeables are the financial prices adjusted by the standard conversion factor 0.85. * The total for
average crop output excludes pumpkins.
Biodiversity and Economics
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423
2. Uses and Values of Biodiversity
Table 27.17 Net present value of benefits from the Hadejia-Jama‘are floodplain,
Nigeria
BASE CASE (8%, 50 YEARS)
TOTAL 1,360 1,
Agriculture 921
Fishing 300
Fuelwood 139
Adjusted agriculture 838
Adjusted total 1,276 1
(8%, 30 YEARS)
251
848
276
127
773
,176
(12%, 50 YEARS) (12%, 30 YEARS)
922 895
625 607
203 197
94 91
574 558
872 846
Source: Barbier, E.B., Adams, W.M. and Kimmage, E. 1991. Economic Valuation of Wetland Benefits: the Hadejia-Jama’are. floodplain, Nigeria.
Notes: values in Naire per hectare; N7.5 = US$1 (1989-90)
Fuelwood production provides an important source of both
tural income and domestic inputs. With the estimated
86,000 rural households in the region each consuming an
average of 50kg of fuelwood per month, total annual rural
fuelwood consumption is approximately 51,600 tonnes
annually.
However these estimated benefits accrue over the lifetime
of the wetlands. The flow of benefits over time from the
continued existence of the wetlands must be converted to a
single number reflecting their discounted net present value.
The overall calculation is acutely sensitive to the discount
rate employed and the time horizon considered.
Indirect use values
In addition to the fuelwood products, wetland forest
reserves yield other non-timber products that are vital for
the rural household economy in developing countries. In the
Hadejia-Jama’are wetlands leaves are harvested from the
doum palm which can be processed into mats and other
household materials or sold unprocessed. Baobab leaves
provide a staple food source as an ingredient in soups and
stews. Mats and other doum products such as baskets and
rope are sold in regional markets or exported to other
localities. The leaves fetch about N20 per sack. Whilst
many of these products are not directly sold, in theory a
value could be attributed to their consumption, using the
price of available substitutes. Livestock and grazing are also
supported by the Hadejia-Jama’are wetlands. To value the
contribution of these inputs to agricultural production we
could assess the costs of alternative means of providing feed
and shelter for livestock. Gradually a figure for each of the
component parts of total economic value that are explicitly
marketed or that have readily available marketed substitutes
can be developed.
Wetland recreational values: USA
The value of a wetland site for recreational purposes proves
to be a more difficult valuation problem. Attempts to place
a value on recreational services yielded by wetlands have
focused on the travel cost approach and contingent
valuation methods. The travel cost approach uses the cost
of travelling to the site as a surrogate for the value yielded
to the recreational consumer of enjoying access to that site.
One study of a wetlands system in Terrebonne Parish,
Louisiana (Farber, 1988), employed windshield
424
questionnaires to assess the costs of travel of respondents.
The costs of access to the site for the different groups were
aggregated to provide a single estimate of ‘willingness to
pay’ for the site of $3,898 million (Table 27.18).
Table 27.18 Estimating willingness to
pay for wetland recreation
site
METHOD OF ANNUAL PRESENT
VALUATION WTP ($) VALUE ($)*
Full Wage 3.898 72.185
0.6 Full Wage 2.733 50.611
0.3 Full Wage 1.860 34.444
Source: Farber, S. and Costanza R. 1987. The Economic Value of
Wetlands Systems.
Notes: Amounts in million US$; * 8% interest and an annual
population growth of 2.6%
Hedonic pricing provides analysts with another means of
deriving values for qualitative environmental attributes. The
assumption is that land values and property values have
such environmental values governing location, air quality,
proximity to sites of natural beauty etc., capitalised in them.
Some portion of the final value of the good or service
exchanged reflects these values. Many studies have focused
on an array of environmental and aesthetic factors to
attribute values to coastal waterfront sites, capturing part of
the aggregate wetland value. Data for land sales in Virginia
Beach, Virginia, covering the period 1953-1976 were used
to estimate a hedonic price equation. It was found that an
increased level of amenity was reflected in increased values,
and that over time the annual value of the amenity was
rising.
Flood control benefits can be estimated using defensive
expenditures: that is the amount needed to be spent to
mitigate against the effect of degrading the wetlands. One
study of the Charles River Basin in metropolitan Boston,
undertaken by the US Department of the Army,
recommended preservation of 8,422 acres of natural storage
areas in the river basin. The estimated value of flood
damages averted by preserving these wetlands was
approximately $80 per acre per year. The value of an acre
of wetland when the flow of flood control benefits had been
capitalised into a single number was estimated at $1,488
(Corps of Engineers).
Biodiversity and Economics
Table 27.19 Values of waterfront amenity in Virginia Beach, Virginia
ARTIFICIAL CHANNEL
land unit 0.1214ha
land unit 0.3035ha
Year frontage 30.48m frontage 45.72m
1955 182 461
1965 192 473
1975 203 557
1985 216 652
1995 231 760
NATURAL BAY
land unit 0.1214ha
frontage 30.48m
land unit 0.3035ha
frontage 45.72m
466 515
568 763
699 1,064
848 1,402
1,016 1,783
Source: Adapted from Shabman, L. and Bertelsen, M.K. 1979. The use of development value estimates for coastal wetland permit decisions. Land
Economics 55:213-222.
Note: Amounts in million US$
Conclusion
Economic development constitutes one of the major threats
to the world’s wetland systems. Land reclamation, diverting
water for irrigation purposes, damming, forestry, and
industrial development are all viewed as the necessary
consequences of advancement and technical change. The
purpose of valuation is to provide a coherent theoretical
basis to examine the costs and benefits of wetland
conversion, in order that the same principles as those
governing other types of investment decisions be adhered
to. Attempts to value wetlands are essentially circumscribed
by knowledge of the functions that the wetlands perform.
This understanding, and thus the associated values may as
yet be imperfect, but are an attempt to ensure that some
values are incorporated into the decision-making process
where perhaps none were before.
PRESERVING FUTURE OPTIONS
Introduction
It is widely argued that a major value of the conservation of
diverse resources arises from their potential contribution to
future mainline production techniques. At some time in the
future it might be useful to include part of the diversity that
now exists in the specialised processes of production. It
follows from this that some amount of variety is then
desirable for the ‘options’ that it represents.
However, studies indicate that valuing future options is not
straightforward. A large amount of theoretical literature and
a small number of empirical studies have examined the role
of uncertainty in valuing uses of natural resources. Option
value (Weisbrod, 1964) reflects the willingness of a risk-
averse society to pay a premium, on top of the use value
itself, for guaranteeing access to a resource of uncertain
future supply. A related concept, often called quasi-option
value (Arrow and Fisher, 1974), is based on information
and the irreversibility of much of the degradation of
biological resources. Quasi-option value attempts to evaluate
the extent to which irreversible changes in natural resources
deny us the opportunity to use future information indicating
new and valuable resource applications.
The intuitive appeal of the concept of option value often
leads conservationists to stress the economic importance of
425
‘preserving options’ by maintaining biodiversity. However,
it should be emphasised that not only has little empirical
work been carried out confirming the magnitude of the
range of possible option values but in theory both option
and quasi-option values may be negative as well as positive.
If future demand for a resource is uncertain, a discount
instead of a premium may be applied to the use value.
Indeed, future demand may become more uncertain as time
passes. Future discoveries may even provide substitutes for
biological resources rather than indicating additional,
profitable uses, leading quasi-option value to be negative.
Randall (1991) suggests that by the early 1980s the
theoretical debate and empirical results had led to a
consensus amongst economists that the sign of option value
is generally indeterminate and that there is little reason to
expect option values necessarily to be of substantial
magnitude.
Nevertheless, there remain persuasive arguments for
believing that the potential value of diversity can be very
large.
First, it is very likely that our current menu of production
does not include the most useful varieties, even under
existing conditions. Active screening programmes have
identified species such as the Rosy Periwinkle which has
yielded great benefit in the treatment of previously
untreatable cancers, including Hodgkin’s disease and
leukaemia. Similarly, close wild relatives of tomatoes and
maize, with extremely valuable characteristics, have been
found in Latin America.
Moreover, because the information and technology that we
have is constantly evolving, there is good reason to expect
that the future usefulness of a particular plant or animal
may be very different from that now recognised. The
rediscovered importance of a dwarf variety of rice provides
a concrete example. As the rice grains were developed for
increased mass, the strength of the stalk became important
(wind could cause all of the nearly mature plants to collapse
making harvest difficult). A dwarf variety of rice with a
short but strong stalk was rediscovered and used in crop
improvement; if this rice variety had earlier been lost there
would have been no way to meet the unforeseen need.
Thus, the conservation of biodiversity provides insurance
against new needs that arise with changing conditions.
2. Uses and Values of Biodiversity
Changing conditions arise for more fundamental reasons
than technological advance. Perhaps the most important of
these are climatic changes and the constant evolution and
changes in distribution of pest organisms and pathogens. In
essence, the movement towards monocultural production
results in a high proportion of the agricultural product being
vulnerable to a single pest. The consequence can be the
collapse of significant portions of the crop. Many of the
crop failures of recent times have been the result of this
interaction between pest and monoculture. Against this
unstable background, it is difficult to know which particular
varieties will be most useful in the long run.
THE VALUE OF DIVERSITY IN PROVIDING
INSURANCE: CROP YIELDS
There will always be a trade-off inherent in specialisation
in production methods. Specialisation implies increased
productivity, manifest in increased average yield, but it also
implies a decreased range of productive assets. Maximum
security is obtained from having the widest possible range
of productive assets; this is known in economics as ‘the
portfolio effect’. It is the basic reason why people tend to
hold their assets in a variety of different forms (e.g. stocks,
bonds, gold and cash). It provides a hedge against the
numerous different risks relating to any one form of asset.
Holding our biological assets in the widest possible variety
of forms would provide this portfolio effect but, potentially,
at the cost of reduced average productivities.
The last 20 years have seen a dramatic rise in international
food-grain production as a result of specialisation, involving
development of high-yield crop varieties, higher inputs,
intensive cultivation and more homogenised farming
techniques. However, as agricultural output has risen, so
has its variability. This increased variance concerns
farmers, governments and policy-makers alike. The
instability of agricultural output may give rise to famines,
regional shortfalls in agricultural produce and at the very
least often results in income instability. The costs of this
variability are not insignificant.
Some theorists argue that these costs are the inevitable
consequence of concentrating the genetic base of many
crops through hybridisation, and that the costs of measures
to reduce the effects of yield variability provide a natural
measure of the costs of loss of biodiversity. The Green
Revolution changed the fortunes of the developing and
developed worlds, helping to alleviate the predicted famines
of the early 1960s and 1970s. It may also provide a key to
developing a value for biodiversity.
The Green Revolution
The ‘Green Revolution’ refers to the rapid increase in
wheat and rice yields in developing countries, brought about
by the use of improved seed varieties and the application of
fertilizers and other chemical inputs. These have made high-
yielding crop varieties comparatively more profitable than
other traditional grains and vegetables and as a consequence
the area sown with improved seed has increased
dramatically.
426
The success of the high-yield varieties is indicated by the
speed at which they have spread across the developing
world. It has been estimated that between one-third and a
half of the area devoted to rice in the developing countries
is now sown with them. CIAT (International Centre for
Tropical Agriculture, Colombia) estimated, for example,
that in the mid-1980s high-yielding varieties were grown on
90% of the 3 million ha devoted to rice in Latin America.
Table 27.20 indicates the area devoted to modern rice
varieties in 11 Asian countries.
There can be no doubt that the Green Revolution has
worked miracles in improving food production in many
parts of the world. Again, CIAT (1981) estimates that yield
increases are between one ton/ha on irrigated areas and
0.75 tons/ha on upland rice areas. This constitutes an
annual increase of about 2.75 million tons of rice, which at
an average price of $200/ton is an increase in the value of
production of approximately $550 million.
Variability of world cereal production
Whilst the beneficial impact of the Green Revolution has
been a greatly increased volume of food-grain production
since the 1960s, one of the hidden costs has been a
simultaneous increase in production variability. While world
cereal production grew at an average yearly rate of 2.7%
between 1960 and 1983, the coefficient of variation (a
measure of variability) increased from 0.028 during the
period 1960-71 to 0.034 in the period 1971-83. This
increased variability appears to result from reduced
diversity in the varieties and practices used in food
production.
In the absence of explicit stabilisation policies, large
fluctuations in agricultural output can feed through into
extreme price variability. Small farmers and the very poor
are particularly vulnerable to such price movements. The
degree of price instability induced can be substantial in
countries with a large agricultural base.
There are two major components of the increase in the
variability of world cereal production:
© increased yield variances (the year-on-year variability of
production from the same field increases);
increased correlations between the yields of different
crops and countries (there is less regional and global
variety available to average out the effects of local
variability).
The second factor is usually the more important of the two:
the loss of diversity is having its greatest impact by reason
of the loss of the ‘insurance’ role that such variety can
provide on a regional basis. For example, comparing yields
for pre- and post-Green Revolution India reveals that
increased variances in grain yields within crops accounts for
less than 10% of the increase in the variance of India’s total
cereal production. The factor that contributed most to
variations in aggregate agricultural output were increased
synchrony in output between regions. Prior to the Green
Revolution, the pattern of agricultural output had been more
Biodiversity and Economics
Table 27.20 Area devoted to modern rice varieties in 11 Asian countries
COUNTRY YEAR
Bangladesh 1981
India 1980
Indonesia 1980
Korea, Rep 1981
Malaysia W 1977
Myanmar 1980
Nepal 1981
Pakistan 1978
Philippines 1980
Sri Lanka 1980
Thailand 1979
1000ha % OF RICE AREA
2,325 22
18,495 47
5,416 60
321 26
316 44
1,502 29
326 26
1,015 50
2,710 78
612 71
80 09
Source: Hazell, P.B.R. 1985. The impact of the Green Revolution and the prospects for the future. Food Reviews International 1(1).
Table 27.21 Extent of genetic uniformity in selected crops
CROP COUNTRY NUMBER OF VARIETIES
Rice Sri Lanka
Rice India
than 10 varieties
Rice Bangladesh
Rice Indonesia
Wheat USA 50% of crop in 9 varieties
Potato USA 75% of crop in 4 varieties
Cotton USA 50% of crop in 3 varieties
Soybeans USA 50% of crop in 6 varieties
diverse, with many different regions cultivating more
diverse crops by more diverse methods and faring
differently according to regional weather and disease
outbreaks. However many of the techniques and crops have
now been standardised, and thus different regions follow
similar output responses. The result is that yields now have
a strong tendency to move up or down together over large
areas of India (Hazell, 1984). This phenomenon is not
exclusive to the developing world. Maize yields in the USA
exhibit a similar, though more pronounced, trend. The rate
of grain yield increased dramatically in the mid-1950s,
rising from about 57kg/ha/yr in the period 1930-1955 to
approximately 133kg/ha/yr for 1955-1985. The variation
around this rising mean has also increased: the coefficient
of variation for 1950-66 is 0.06, but 0.105 for 1967-85.
The other source of increased variability is the common
genetic base of the different crop varieties. The existence of
genetic variety within the species itself provides insurance
in the same way as variety in crop and cultivation practice
within a nation. With increasing genetic uniformity at the
species level (Table 27.21), production over a wide area
become susceptible to a single external impact, such as a
particular pest or disease.
For example, in 1970 a particular form of corn leaf blight
(Helminthosporium maydis) struck in Florida, spreading
northwards throughout the corn belt, drastically reducing
From 2,000 varieties in 1959 to 5 major varieties today
75% of varieties descended from one maternal parent
62% of varieties descended from one maternal parent
74% of varieties descended from one maternal parent
SOURCE
Rhoades, 1991
Hargrove et a/., 1988
From 30,000 varieties to 75% of production from less
Rhoades, 1991
Hargrove et a/., 1988
Hargrove et a/., 1988
NAS, 1972
NAS, 1972
NAS, 1972
NAS, 1972
yields. Only certain types of hybrids were known to be
susceptible to this blight. Unfortunately, a large proportion
of maize growers in the USA were cultivating such
varieties. Competition over yields and quality had led
farmers to concentrate their dependence on a few plant
varieties with a narrow genetic base. Table 27.22
demonstrates that this phenomenon has been associated with
numerous large-scale crop failures.
Concentrating the genetic base: the case of rice
The widespread adoption of a relatively small number of
improved rice varieties, many of which are closely related
genetically, has gradually reduced the genetic diversity of
the crop. It has been estimated that about 40% of the
world’s rice crops comprise high-yielding varieties. One
variety introduced into Asia in the late 1960s was IR8. This
had a high yield potential with short stiff straw that allowed
it to produce heavy panicles of grain without falling over.
IR8 was also insensitive to photoperiod (daylight hours) or
growth duration, which meant that it could be grown at any
time of the year. IR8 and other semi-dwarf rice varieties
were rapidly adopted throughout Asia, with dramatic
results. Average rice yields for Asia in the period 1971-
1980 were 42% higher than in 1951-1960. Total production
rose by 77% at a time when the land area devoted to rice
cultivation rose by only 25%.
2. Uses and Values of Biodiversity
Table 27.22 Past crop failures attributed to genetic uniformity
DATE LOCATION CROP CAUSE AND RESULT SOURCE
900 Central Maize Anthropologists speculate that the collapse of the Classic Rhoades, 1991
America Mayan Civilization might have been a result of a maize virus
1846 Ireland Potato Potato blight led to famine in which 1 million died and Hoyt, 1988
1.5 million emigrated from their homeland
late 1800s = Sri Lanka Coffee Fungus wiped out homogenous coffee plantations on the island Rhoades, 1991
1940s USA US crops lost to insects has doubled since the 1940s Plucknett and Smith, 1986
1943 India Rice Brown spot disease aggravated by typhoon destroyed crop Hoyt, 1988
starting the ‘Great Bengal Famine.’
1953-54 USA Wheat Wheat stem rust affected most of hard wheat crop Hoyt, 1988
1960s USA Wheat Stripe rust reached epidemic proportions in Pacific Northwest Oldfield, 1984
1970 USA Maize Decrease in yield of 15%, $1 billion lost* NAS 1972, Tatum, 1971
1970 Philippines Rice HYV rice attacked by leafhoppers spreading tungro virus Hoyt, 1988
& Indonesia
1972 USSR Wheat Crop badly affected by weather Plucknett et a/, 1987
1974-77 Indonesia Rice Grassy stunt virus destroyed over 3 million tonnes of rice Hoyt, 1988
: - from the late 1960s to the late 1970s the virus plagued
South and Southeast Asian rice production
1984 Florida Citrus Bacterial disease caused 135 nurseries to destroy 18 million trees Rhoades, 1991
Notes: * Duvick (1986) reports that although the leaf blight attacked a widespread and uniform genotype, the problem was uniformity of cytoplasm -
introduced to eliminate the chore of detasseling - not the genetic material in the nucleus of the seed.
High yielding varieties have had similar successes elsewhere
in the developing world. Some theorists stress that no
conclusive evidence has yet been found that a common
ancestry might contribute to production variability and it
could be argued that certain modern varieties (such as IR64)
have a very diverse parentage and should perform well
under a wide range of conditions. However, it is
increasingly apparent that some varieties share many genetic
elements. The genetic parentage of IR8 can be traced to
1914, when a variety of rice called Cina was introduced
from China into Indonesia where it attained rapid popularity
because of its photoperiod insensitivity, its high yields and
grain quality. In 1934 plant breeders in Indonesia crossed
Cina and Latisail (an Indian variety) to form Peta. In 1962
Peta was used as the female parent in a cross with Dee-geo-
woo-gen, a semi-dwarf stiff-strawed rice variety from
China. IR8 was one of the progeny from this cross. In the
1970s and early 1980s further hybridisation using IR8 and
Peta derivatives as parents produced many of the semi-
dwarf varieties that are now grown worldwide. Components
of the cytoplasm (the protoplasm of a cell excluding its
nucleus) are inherited through the female parent, so
varieties with Cina as their ultimate maternal ancestor
probably carry similar cytoplasm. In 1983-1984, 38% of a
sample of the female parents used in 106 crosses were the
maternal progeny of Cina, which implies that many of the
varieties selected from these crosses and released in the late
1980s will also carry similar cytoplasm.
The rice plant is most vulnerable to stress during its
reproductive growth phase. Photoperiod-sensitive varieties
grown traditionally in tropical Asia entered the reproductive
phase during the peak rainfall period when risks were
minimised. These varieties then ripened at the end of the
rainy season. With the introduction of IR8 and other similar
varieties with photoperiod insensitivity, farmers in many
428
latitudes were able to plant and cultivate at any time of the
year. Those in irrigated areas were also able to plant
several rice crops annually instead of one, as previously.
Varieties that mature a fixed period after seeding are often
more vulnerable to climatic changes and natural disasters
such as typhoons or droughts. This can be offset by
strategic staggered planting, which would ensure staggered
maturity. However, because planting is a particularly
labour-intensive activity, staggered planting would require
radical changes in the agricultural labour market (from
seasonal labour flows to continual rolling employment).
Various institutional changes would have to come about to
ensure this, and until such changes occur the fixed growth
duration varieties may have less ‘buffering capacity’ to
withstand extreme climatic variation than the traditional
photoperiod-sensitive varieties.
Reduced plant height is one of the more obvious
characteristics of modern rice plant varieties. This improves
the harvest index and the ratio of grain to straw, and allows
the plant to remain standing after heavy doses of nitrogen
fertilizer. It is this characteristic that has been hailed as the
one most responsible for the production yield gains of the
Green Revolution. Unfortunately drought during the
vegetative growth stage can shorten the height of semi-
dwarf varieties to significantly below optimal levels. This
undermines their ability to withstand subsequent floods and
weed growth. Although most modern rice varieties continue
to be semi-dwarfs there may be a gradual shift towards
taller varieties. In 1975 69% of new varieties were
semi-dwarf with a mean height of less than 130cm whereas
in 1984 54% of new varieties were semi-dwarf.
Traditional rice varieties were naturally selected over a
period of centuries for their resistance to or tolerance for
their local environments. Modern rice varieties are the
product of less than a century of genetic experimentation,
with many varieties being selected in less than a decade
(IR36 was developed in five years). The experimental sites
have often been radically altered by the application of
pesticides and fertilizers. As a result of this
experimentation, pest-resistant varieties have been
developed with inbuilt responses to many common rice
pests for which they have been specifically screened.
However, it seems unlikely that modern varieties could
have generic resistance comparable to that of traditional
types. It is even feasible that pest attacks are more
concentrated or widespread because extensive areas are
being sown with more homogenous crops.
Drought tolerance is not generally a feature of modern rice
varieties. Modern rice plants are bred to produce the
maximum amount of grain under optimal conditions. In the
process of genetic refinement, many of these varieties have
lost the deep root system required for drought tolerance.
IR5S2 has been developed in response to this loss in root
depth, but the apparent yield potential of such varieties is
significantly lower than that of their less drought-tolerant
counterparts.
Similar case histories can be related for wheat and maize
which emphasise that, whilst a common genetic base is not
necessarily cited as the sole cause of increased yield
variability, it may play an important part in causing
co-movements in grain yields world wide.
SOURCES OF YIELD VARIABILITY
Tables 27.23 and 27.24 illustrate the main components of
variability in world agricultural cereal production. These
figures reveal that increases in mean yields account for
about 70% of the increase in total cereal production and
expansion in area for 20%, and also that wheat and maize
contribute greatly to the change in mean total cereal
production (32.65% and 35.18% respectively). Table 27.24
indicates the percentage change in the variance of world
cereal production attributed to its components. The column
sums show that 95.93% of the increase in the variance of
world cereal production is attributable to changes in the
Biodiversity and Economics
variances and covariances of crop yields. The change in
maize yield variances and covariances accounts for 17.16%
of the overall increase in the variance of world cereal
production. Changes in area-yield covariances exerted an
important stabilising effect on world cereal production,
reducing the variance of total cereal production by 42.28%.
Virtually all of this reduction can be attributed to a decline
in area-yield correlations, of which the strongest declines
appear to be between crop yields in one country and the
sown areas of different crops in different countries 28.51%.
Genetic uniformity and crop yield variability
Genetic uniformity has been cited as one of the major
causes of widespread yield reductions in maize in 1970 in
the USA. At that time approximately 80% of US maize was
based on T cytoplasm, which is particularly susceptible to
the T race of southern corn leaf blight (Tatum, 1971). The
rapid spread of this fungus across the eastern part of the
nation was aided by an abnormally wet summer which
increased the germination and dissemination of its spores.
Hybrids without the T cytoplasm were unaffected.
The direct costs of genetic uniformity and of monocultural
production have been an increase in the vulnerability of
crops and regions to climatic variations and to disease. The
fact that common wheat and rice varieties such as Bezostaia
wheat in Eastern Europe and IR36 rice in Asia have been
cultivated extensively (more than 10 million ha in each
case), increases the risk of crop failure in the event of an
epidemic.
Technical uniformity: inputs and crop yield variability
With the international adoption of genetically engineered
seed types there has been a worldwide revision in
cultivation techniques. The application of chemical inputs
has dramatically increased over the last three decades.
Many argue that one of the main causes of agricultural
output variability, and especially of grains and cereals, is
varying levels of input use in response to price and interest
rate movements.
Table 27.23 Components of change in world average cereal production 1960-1971
to 1971-1983
COMPONENTS OF WHEAT MAIZE RICE BARLEY MILLET SORGHUM OATS OTHER TOTAL
CHANGE
% change in mean yields 80.93 64.21 60.62 39.52 63.64 45.63 -528.08 -179.99 72.40
% change in mean areas 14.94 28.61 33.64 49.11 44.76 44.42 534.84 220.53 22.36
Change in area-yield 0.19 0.09 -0.02 0.45 2.96 0.20 15.21 -1.08 00.14
covariances
Contribution of crop to 32.65 35.18 11.50 18.28 0.55 4.34 -0.47 -2.03 100.00
change in mean production
of total cereals
Source: Hazell, P.B.R. 1989. Changing patterns of variability in world cereal production. In: Anderson, J. and Hazell, P. (Eds), Variability in
Grain Yields, Implications for Agricultural Research and Policy in Developing Countries.
Note: In per cent; excluding China.
429
2. Uses and Values of Biodiversity
Table 27.24 Components of change in the variance of world cereal production
1960-1971 to 1971-1983
CHANGE IN CHANGE IN
MEAN YIELDS MEAN AREAS
Crop variances
Wheat 2.06 -2.38
Maize 6.67 1.94
Rice 0.11 0.25
Barley 0.43 2.30
Millet 0.01 -0.01
Sorghum 0.19 0.07
Oats 0.83 0.27
Other 0.14 -0.15
Sum Crop Variances
within Countries 10.44 2.28
Intercrop Covariances
within Countries 0.97 4.48
Intercountry Variances
within Crops 0.09 1.61
Covariances between
different crops in
different countries 2.75 0.85
Column sums 14.24 gr22
SOURCE OF CHANGE
CHANGE IN YIELD CHANGE IN AREA CHANGE IN
VARIANCES AND VARIANCES AND AREA-YIELD
COVARIANCES COVARIANCES COVARIANCES
5.27 -0.57 3.57
17.16 -6.15 -5.01
0.45 0.12 0.16
1.87 0.86 1.37
0.04 0.01 0.06
0.57 -0.23 0.12
0.11 -1.25 -0.54
0.93 -0.14 0.29
26.40 -7.36 0.01
36.68 -0.94 -9.38
11.49 -3.61 -4.40
21.36 19.13 -28.51
95.93 7.22 -42.28
Source: Hazell, P.B.R. 1989. Changing patterns of variability in world cereal production. In: Anderson, J. and Hazell, P. (Eds), Variability in
Grain Yields, Implications for Agricultural Research and Policy in Developing Countries.
Most of the studies in Table 27.25 support the view that
increased application of nitrogen increases variance in yield.
In most cases, the change in variance with respect to
nitrogen is higher than change in mean yield. According to
Byerlee and Anderson (1969), with a nitrogen level of 20
kg/ha, a 1% increase in the nitrogen level would result in
a 0.08% increase in the mean yield and a 0.44% increase
in the variance of output. Where the nitrogen level is 40
kg/ha, a 1% increase in nitrogen level results in a 0.04%
increase in the mean yield but a 0.62% increase in the
variance of output. The supply of other inputs such as
irrigation or the application of pesticides and herbicides may
also affect the variability of crop yields. How the
application of such inputs affects yields and their variance
is not yet fully recognised.
CROP INSURANCE: THE RESPONSE TO
INCREASED AGRICULTURAL RISK
One possible response to increased yield instability and
consequent income variability is to acquire insurance.
Insurance schemes typically offer a means of guaranteeing
expected future income in the face of uncertainty. This is
accomplished through the payment of a premium which
ensures that an indemnity is received in the event of an
undesirable outcome. The amount of this indemnity is
usually sufficient to compensate the individual for the loss.
In other words, the essence of an insurance programme is
the sacrifice of some amount (the ‘premium’) of the average
return from the activity in return for a reduction in the
long-term variability of returns.
This is one of the major roles of diversity in agricultural
production. Reduction in variety of species and techniques
430
has raised average returns but also increased variability.
Conversely, increases in diversity in agriculture provide
insurance, by reducing variability in return for a reduced
mean return. Biodiversity can be said to provide a form of
natural insurance.
One means of reducing the risks associated with yield
fluctuations is to diversify the portfolio of crops, moving
away from monoculture. Intercropping, spatial
diversification, staggered planting, and hoarding are
surprisingly efficient in reducing income risks. Such
practices have been employed by agriculturalists for
centuries.
However, diversity is not the only means of providing crop
insurance. The market itself will do so, if the risks are
insurable. There is not necessarily any reason to intervene
if this is the case, because farmers themselves could then
choose the least expensive basis for insuring their crops,
allocating their ‘insurance policies’ between the market and
diversity.
However, this is only the case if insurance markets are able
to operate effectively. In many cases they do not. This is
because market insurance operates by means of the pooling
of independent risks. That is, in many circumstances
individuals may face uncertainty, but society as a collective
of individuals faces approximate certainty. This is
attributable to ‘the law of large numbers’. In essence,
insurance works effectively when an individual farmer does
not know whether his/her crops will fail this year, even
though the failure rate for crops in that region for any given
year is known and relatively stable over time.
Biodiversity and Economics
Table 27.25 Changes in mean and variance of crop yield with respect to nitrogen
fertilizer
STUDY/SOURCE CROP
Anderson, 1973 Wheat, Australia
Smith and Umali, 1985
Antle and Crissman, 1986
Rainfed rice, Philippines normal*
Rainfed rice, Philippines gamma”
Rice, Philippines 1975-76*
1977-79*
Byerlee and Anderson, 1969 Wheat, Australia
Ryan and Perrin, 1973 Potatoes, Peru
Roumasset, 1974 Rice, Philippines
Village 1
Village 2
Village 3
Rosegrant and Herdt,
1981
Rice, Philippines
Irrigated
Rainfed
Smith et a/., 1984 Rainfed rice, Philippines
Wet Season
Dry Season
Rosegrant and Roumasset,
1985
Rice, Philippines
Average irrigation, dry season
Average irrigation, wet season
Rainfed, wet season
Good irrigation, dry season
NITROGEN MEAN VARIANCE
LEVEL (Kg/ha)
40 0.14 0.22
80 0.06 0.19
40 0.28 0.36
80 0.16 -0.08
11 0.16 0.22
21 0.25 -0.35
20 0.08 0.44
40 0.04 0.62
100 0.10 0.20
200 0.17 0.35
40 0.49 0.49
80 0.16 0.32
40 0.22 0.45
80 0.03 0.06
40 0.19 0.37
80 0.00 -0.01
40 0.20 0.29
80 0.19 0.42
40 0.14 0.31
80 0.10 0.30
40 0.16 0.24
80 0.10 0.48
40 0.15 0.26
80 0.10 0.48
40 0.20 0.03
80 0.24 0.36
40 0.19 0.12
80 0.21 0.54
40 0.14 0.06
80 0.13 0.49
40 0.13 0.14
80 0.10 0.59
Source: Adapted from Roumasset, J.A., Rosegrant, M.W., Chakravorty U.N. and Anderson J.R. 1989. In: Anderson, J.R. and Hazell, P.B.R.
(Eds), Variability in Grain Yields, Implications for Agricultural Research and Policy in Developing Countries.
Notes: * Reported elasticities are computed at mean input levels, expressed in Pesos per hectare. Figures given are estimated mean nitrogen use
given prevailing prices. * Yield distribution is assumed to be normal or gamma as specified.
The primary assumption that drives the insurance principle
is that the probability of a crop failure for any given
individual is independent of that for anyone else. That is,
when risks are faced by all persons uniformly, it is not
possible for an insurance market to operate. This is because
it does no good to ‘pool’ a risk if everyone will incur the
loss at the same time.
It is apparent that the assumption of independence fails in
the case of crop insurance in the USA. The agricultural
sector is one that faces pronounced co-movements in output.
It is self-evident that individual agents’ probabilities of
experiencing a crop failure are not independent when
techniques and varieties become standardised. This is borne
out heavily in the data. The government is required to
subsidise the insurance companies in order that continued
cover can be provided.
In short, the crop insurance market in the USA has not
operated effectively, probably on account of the correlation
431
of risks. The US experience demonstrates the difficulty in
developing and administering crop insurance cover, with the
private sector being unwilling to provide complete
insurance. The current insurance programme dates only to
the Crop Insurance Act of 1980, which allowed private
insurance schemes to operate in this area, but the evidence
from this period is clear. The Federal Crop Insurance
Corporation (FCIC) currently subsidises the premiums paid
by farmers by about 30%. The amount of government
subsidy can be seen in the difference between Total
Premium and Farmer Premium (Table 27.26). The total
costs of the protection offered including the subsidy and
administration costs are shown in Table 27.27.
During the 1980s, the US government spent $3.8 billion on
crop insurance programmes for US farmers. This is very
important for two reasons. First, it is indicative of the
extent of crop failures occurring under specialised
agriculture. Second, and more important, it is obvious that
these markets were requiring substantial government
2. Uses and Values of Biodiversity
Table 27.26 Summary’ of multiple peril crop insurance protection in USA
YEAR PROTECTION TOTAL ACRES TOTAL
INSURED PREMIUM
Million $ Thousands Million $
Total ‘81-90 75,592 753,468 4,751
1981 5,981 58,324 377
1982 6,125 54,918 396
1983 4,370 36,542 286
1984 6,620 55,492 434
1985 7,167 63,360 440
1986 6,219 64,004 380
1987 6,079 64,794 365
1988 6,957 73,799 436
1989 13,563 139,365 816
1990 12,511 142,870 821
Source: American Association of Crop Insurers (1991).
Note: ' Summary of all crops for all states by year.
Table 27.27 Nature and extent of all government costs
PREMIUM EXCESS
SUBSIDY LOSSES
Total 684,583 1,731,597
1981 46,995 30,471
1982 91,990 132,250
1983 63,669 297,971
1984 98,296 204,314
1985 100,224 242,438
1986 88,043 233,806
1987 87,536 4,669
1988 107,830 585,678
FARMERS LOSSES Loss FARMERS
PREMIUM PAID RATIO BENEFIT/COST
Million $ Million $ RATIO
3,648 6,912 1.46 1.89
330 407 1.08 1.23
305 529 1.34 1.74
222 584 2.04 2.63
336 638 1.4 1.90
340 683 1.55 2.01
291 613 1.62 2.10
277 369 1.01 1.33
328 1,049 2.41 3.20
610 1,189 1.46 1.95
609 851 1.04 1.40
FCIC MMA REINSURANCE TOTAL
COSTS COSTS * COSTS COSTS
562,356 178,705 560,583 3,717,824
60,630 27,658 3,663 169,417
69,190 46,978 23,138 363,546
69,745 25,958 35,603 492,946
73,632 25,235 78,887 480,364
79,009 17,711 102,888 542,270
85,027 10,765 97,711 515,352
60,046 12,700 97,148 262,099
65,077 11,700 121,545 891,830
Source: Report of the Commission for the Improvement of the Federal Crop Insurance Program. Washington DC.
Note: Figures in thousands US$; * direct agent costs Master Marketers.
subsidies for operation. In the period 1981-1988, the US
government spent $685 million on direct subsidies in order
to encourage the operation of the market.
This crop insurance programme both indicates the value of
diversity and discriminates against it. If diversity can itself
provide insurance against widespread crop failures, then this
value would accrue to practices which maintained diversity.
Although insurance through diversity would not be a policy
operated through the financial markets, it could just as
effectively generate this value as one that does.
being allowed to operate. It is instead being pre-empted by
a government policy that is encouraging, through subsidy,
the substitution of the financial market. This sort of policy
discourages farmers from using natural diversity for the
provision of insurance, even when it is the most effective
means of doing so (Swanson, 1992).
THE VALUE OF AGRICULTURAL GENETIC
DIVERSITY
One area in which the actual value of qualitative diversity
has been estimated is agricultural genetic diversity. Here,
the closest relatives to the small number of domesticated
species are often investigated to ascertain their potential for
contributing to the productivity or resilience of the domestic
variety.
Yield gains in agriculture are typically broken down into a
technology component (encompassing chemicals and capital
machinery) and a genetic component. Gains from crop
breeding arise from genetic improvements in a number of
different fashions:
the environmental conditioning of the plant (e.g. better
standibility, drought resistance, etc.)
pest and disease resistance
suitability to changing cultivation technology (e.g.
response to fertilizers)
more productive genotypes (e.g. number or size of
kernels)
quality characteristics (e.g. changes in protein or oil
content).
A considerable amount of work has been carried out in
estimating the often substantial value of genetic
improvements to crops. Some of the more important
studies are summarised in Table 27.28.
The aggregate value of the raw genetic materials used in
crop-breeding is best ascertained by reference to the
industry’s spending on research and development. This is
because, as with so many of the facets of biodiversity, the
value of genetic variety for crop breeding lies in the
potential value of future finds from the existing genetic
Biodiversity and Economics
Table 27.28 Genetic diversity and agriculture: genetic contributions of cultivars to
crop yields
CROP LOCATION PERIOD EFFECT ON PRODUCTION SOURCE
All crops USA 1980s $1.0 billion/year OTA, 1987, USDA est.
Maize USA 1930-80 = % of a fourfold increase in yields OTA, 1987
USA 1930-80 89% of yield gain of 103 kg/ha/yr in commercials Duvick, 1984
USA 1930-80 71% of yield gains in single cross hybrids Duvick, 1984
USA 1985-89 Genetic gains to N. Dakota of $2.3 million/year Frohberg, 1991
Rice Asia GR $1.5 billion/year Walgate, CALP
USA 1930-80 = % of a doubling in yields OTA, 1987
Wheat Asia GR $2.0 billion/year Walgate, CALP
USA 1930-80 = % of a doubling in yields OTA, 1987
USA 1958-80 0.74% genetic gain per year - % of 32% yield gain Schmidt, 1984
UK 1947-75 50% of an 84% gain in yields Silvey, 1978
World 1970-83 43% of genetic gain totalling 46% (best data) Kuhr et a/., 1985
55% of genetic gain totalling 32% (all sites) Kuhr et a/., 1985
Sorghum USA 1930-80 = % of a fourfold increase in yields OTA, 1987
1950-80 1-2% genetic gain per year from manipulating Miller and Kebede, 1984
kernel numbers, plant weight, height and leaf area
Barley USA 1930-80 = % of a doubling in yields OTA, 1987
Potato USA 1930-80 = % of a fourfold increase in yields OTA, 1987
Soybeans USA 1930-80 = % of a doubling in yields OTA, 1987
USA 1902-77 79% of 23.7 kg/ha annual yield gains Specht and Williams, 1984
Pearl Millet India at present genetic improvements worth $200 million annually ICRISAT, 1990
Cotton USA 1930-80 = ‘4 of a doubling in yields OTA, 1987
1910-80 0.75% genetic gain per year Meredith, Jr and Bridge, 1984
Sugar cane USA 1930-80 = ‘% of a doubling in yields OTA, 1987
Tomato USA 1930-80 = % of a threefold increase in yield OTA, 1987
Table 27.29 Genetic diversity and agriculture: specific contributions made by wild
relatives of crops
CROP FOUND IN EFFECT ON PRODUCTION SOURCE
Wheat Turkey Genetic resistance to disease valued at $50 million per year Witt, 1985
Rice India Wild strain proved resistant to the grassy stunt virus
Barley Ethiopia Protects California’s $160 million per year crop Witt, 1985
from yellow dwarf virus
Hops Added $15 million to British brewing industry in 1981 by Witt, 1985
improving bitterness
Beans Mexico The International Center for Tropical Agriculture in Colombia used Rhoades, 1991
genes from the Mexican bean to beat the Mexican bean weevil
which destroys as much as 25% of stored beans in Africa and
15% in South America
Grapes Texas Texas rootstock (from land now covered by the Rhoades, 1991
Dallas-Fort Worth Airport) was used to revitalise the
European wine industry in the 1860s after a louse infection
stock. An indication of this value is given by the returns
Tealised from past efforts at developing the previously
existing gene pool for commercial use, as well as by the
amounts currently being invested in such efforts.
The top 25 agricultural biotechnology - or crop breeding -
firms spent $330 million on research and development in
1988 (Hobbelink, 1991). Crop breeding has generated a
433
large return in the past - US public and private expenditures
on corn research totalled $100 million in 1984 contrasted
with an estimated return of $190 million (Huffman and
Evenson, 1991). These figures both indicate that there is
considerable value to be had from retaining substantial
variety in the plants that are most closely related to our
domesticated crops. Several important examples are given
in Table 27.29. These varieties represent only a fraction of
2. Uses and Values of Biodiversity
existing biological diversity, but are probably some of the
most valuable species to retain on account of the ease of
their introduction into mass production.
The calculated value-gains from crop-breeding efforts are
not, however, equivalent to the value of the raw genetic
material that exists in the wild, for two main reasons. First,
such gains may be achieved using raw materials from a
variety of sources: existing cultivated varieties (cultivars),
varieties husbanded by traditional farmers (land races), wild
relatives of crops or even - with the advent of genetic
engineering - completely unrelated species. Second, these
gains must be apportioned amongst a number of factors
which, together with these raw genetic materials, generate
this increased value, including scientific effort, technology
and commercial development.
THE VALUE OF BIODIVERSITY IN THE
PRODUCTION OF PHARMACEUTICALS
The medicinal value of plants and their derivatives has been
recognised for millennia (see Chapter 25). Estimating the
importance and economic value of the biodiversity which
gives rise to the possibility of more discoveries is a very
recent field of interest.
The basis of much of the estimation is a very detailed
survey which was carried out on those prescription drugs
(in the USA) which were derived in some way from
flowering plants (Farnsworth and Soejarto, 1985). The
study involved determining the basic materials in all of the
thousands of different drugs prescribed in the USA over the
period 1959 to 1973 and then identifying those which were
plant-based (see Table 25.6 for examples). This was taken
to include those drugs which contained crude plant extracts,
semi-purified mixtures of active principles, single active
principles or active principles which had been chemically
modified.
It was found that, for the period examined, the proportion
of plant-based drugs was just over 25% of all prescription
drugs (in a market where the 1973 value of the total
prescription drugs sales was over $6.3 billion at retail
prices). On this basis the value of plant-based prescription
drugs was estimated to be about $1.6 billion in 1973 and
the additional value of the same drugs provided directly
through hospitals and clinics was probably as much again,
giving a total value of about $3.2 billion.
The authors also estimated a figure for 1980 on the same
basis and obtained a total of around $8.2 billion (in current
prices). A later study (Principe, 1991) using a variation of
this approach but including an estimate of non-prescription
drugs revised the 1980 figure to $9.8 billion and calculated
a 1985 value of $18 billion (all of these figures being for
US sales alone).
Interestingly, the pharmaceutical industry use of plant
diversity has been dependent upon a small number of
species. The authors of the first study found that, of the
25% of pharmaceuticals traceable to plant-based origins, a
mere 40 species of plants were at the ultimate source. Using
their figure of total retail value of $8 billion gives an
average value per species utilised of $200 million, though
of course there is a large amount of variability.
434
These figures give an indication of the direct retail value of
plant-based materials in medicine. The numbers are very
large and can probably be trebled to give a worldwide total
because the US market represents about one-third of world
pharmaceutical sales.
However, it must be remembered that these values are retail
market figures, and not only the value of the plant material
on which the drugs are based. The price of the raw
materials themselves may be of the order of only a few per
cent of the final market value but their economic value to
the drug industry is far more than their basic cost.
Estimation of the real economic value is a conceptual
problem as much as a practical one and is discussed below.
The value of the underlying biodiversity which has
generated these plant-based drugs and which may give rise
to many others is an even more difficult issue.
With successful plant-based drugs having a very high
potential value it might be expected that the pharmaceutical
industry would be very active in research in this area but
the industry’s attitude appears to be somewhat ambivalent.
New drugs are developed through two broad approaches:
the screening of potentially active material for medical
usefulness and/or the synthesis of specific types of
compounds based on the understanding of biochemical
reactions within the human body. Recently, many of the
most successful modern drugs have come through the
application of the techniques of biotechnology and genetic
engineering, and there has been a movement away from
lengthy and costly screening processes. Even more recently,
however, there appears to be a resurgence of interest, on a
small scale at least, in screening approaches (Findeisen,
1991) . The reasons for these shifts in emphasis will also be
discussed below.
What role do plants play in pharmaceutical production?
Three major ways have been identified in which plants are
used within the pharmaceutical industry (Principe, 1991).
These are:
constituents isolated from plants are used directly as
therapeutic agents
plant constituents are used as base materials for the
synthesis of useful drugs
natural products are used as models for the synthesis of
pharmacologically active compounds.
The first two of these uses represent market values of
natural plants as raw materials consumed directly in the
pharmaceutical industry. These are the uses which have
been valued in the billions of dollars by the studies cited
above. However, it has already been noted that the raw
material value is usually only a very small proportion of the
overall retail price of the drugs which includes factors such
as store rental, employees’ salaries, transport and taxes.
Therefore, estimates based on retail value necessarily
represent upper-bounds on the raw material values.
There is good reason to believe that the cost of the raw
materials used directly in pharmaceutical manufacturing
must remain low. This is because it is generally possible to
synthesise chemical substances artificially if the costs of the
natural material are too high. Once the method of operation
is identified, the cost of chemical batch processing is
generally very low, and artificial synthesis of the active
ingredients usually becomes the least-cost mode of
production for mass-produced substances. For example,
aspirin is now produced synthetically although the original
source was the bark of the willow tree.
For this reason, it cannot be expected that the direct use of
plant variety for pharmaceutical manufacture will ever be
very substantial, or that it will be possible to claim high
returns for presently unpatentable natural products. For
example, the Mexican government has historically been a
major producer of the yam Dioscorea, which has been the
source of the basic material used in the production of
steroid drugs sold as oral contraceptives and cortisone. This
market was producing nearly $83 million annually for
Mexico in 1976 (Oldfield, 1984). However, as the Mexican
government attempted to extract a higher return from the
export of the yam by raising prices, the pharmaceutical
manufacturers turned to synthetic processes and the market
for Dioscorea collapsed (Principe, 1991). Therefore, given
the ready alternative of artificial synthesis, direct use values
will never be very substantial (there are exceptions to this
general rule, namely: reserpine, codeine, morphine,
digitoxin, and atropine (Oldfield, 1984)).
Despite advances in medical science and progress in
biochemical engineering, there are many conditions and
diseases for which we currently have no effective treatment.
As long as untried or unknown plant species exist so do the
possibilities for discovering materials which could lead to
important new drugs.
A topical example of this is the development of the drug
Taxol and its derivatives. Taxol is a compound obtained
from the bark of the Pacific Yew Taxus brevifolia and has
been demonstrated in clinical trials to be effective in
treating certain difficult ovarian and breast cancers.
Unfortunately the Pacific Yew tree is extremely slow
growing and the bark from several trees would be required
to provide sufficient Taxol to treat one patient. Several lines
of development are being pursued, from high technology
chemical synthesis techniques (which have so far had
meagre success) to the planting of large numbers of yews
in commercial forests. A promising approach is the
isolation of a related but possibly more powerful compound
from the leaves of the same yew tree, leading to the
prospect of harvesting the compound without killing the tree
(Potier, 1991).
This example illustrates the potential for plant products. A
highly promising drug is being developed, based on the
efficacy of a natural compound. The active ingredient is
very difficult to synthesise but research continues on
synthesis and on naturally occurring variations. Whatever
form the final commercial product takes it will have been
derived from the discovery of the properties of the basic
natural compound. Nature has, in effect, provided the
blueprint for a drug which is effective in fighting cancer,
and while biochemical engineers may modify the original
design these are only incremental changes.
Thus, the most important value of plants in this context lies
in the information which they can provide; specifically,
435
Biodiversity and Economics
information about the possible existence (and possible loss)
of natural blueprints for drug design.
What is the value of the information contained in plant and
animal diversity? First, it is the value of the chance
discovery, i.e. one that proceeds from mere trial and error.
One straightforward attempt at such a valuation has been
attempted (Farnsworth and Soejarto, 1985; Principe, 1991).
Its method was to look at the success rate for those plants
that have been surveyed for their pharmaceutical benefits,
assuming those species to be randomly chosen. As earlier
studies had estimated that 5,000 plant species had been
thoroughly examined for medicinal effectiveness, and since
there are 40 species in use in prescription drugs, the
assumption of randomness would suggest that one in 125
randomly selected species would be developed to a
successful product. Thus for every 1,000 species which
becomes extinct, eight potentially useful plant-derived drugs
would be lost. At the average retail value of $200 million,
this would lead to pharmaceutical losses of $1.6 billion in
retail value. In this case, retail value is a useful measure of
the willingness-to-pay for the information which is assumed
to be a prerequisite to the existence of the particular drug.
Consumers demonstrate that they value the existence and
discovery of this information through their willingness to
purchase the drug at its shelf price. However, it should be
stressed that in practice species are not chosen for medical
screening at random but are pre-selected. Therefore among
1,000 plant species chosen at random, there may be
expected to be fewer than eight potentially useful plant-
derived drugs; nevertheless this form of valuation gives a
useful approximation of what may be lost.
This valuation methodology stresses the experimental nature
of pharmaceuticalcompany research. Although this example
requires the use of averages, in fact the pursuit of new
drugs is much more of a lottery than even these numbers
would suggest. If the company’s experiments result in a
major discovery, such as Taxol appears to be, a single drug
can be as valuable as many other entire industries. The
sales and profits of a best seller can be very high: in 1990,
the top selling drug world wide (Zantac - an ulcer medicine)
grossed sales of about $2.4 billion. Nine drugs earned over
$500 million each in the USA alone (which probably
indicates per drug earnings of about $1 billion world wide).
Pharmaceutical companies must reject hundreds if not
thousands of possibilities before one of these discoveries is
unearthed. Nevertheless, this method of research and
discovery is not haphazard, although imbued with chance:
five of the top 20 most profitable companies in the world
are pharmaceutical companies.
This profitability is partially attributable to the fact that
significant discoveries are awarded monopoly rights for a
period of 10-20 years, which generates substantial returns
to the successful experiment. However, this profitability is
attributable to the fact that the search is not entirely
random. The companies utilise all of the information on
chemistry, physiology, and other experimental evidence that
is available in order to guide them.
One very important form of experimental evidence available
to pharmaceutical companies is the experience of peoples
living in contact with plant and animal species. These
2. Uses and Values of Biodiversity
communities have had, in most cases, thousands of years of
trial and error experimentation in order to build a record
regarding plant usefulness. This indigenous knowledge is
the directory which provides the indicator concerning which
species are most useful in terms of chemical effects. With
the use of this knowledge, search by pharmaceutical
companies need not be random.
Recently, a return to greater interest in plant opportunities
and to screening approaches seems to be occurring. In
1988, 17% of total pharmaceutical industry research and
development spending in the USA went on ‘Biological
Screening and Pharmacological Testing’ (Pharmaceutical
Manufacturers Association, 1988-1990). This represents
expenditures of over $1 billion dollars although the amount
actually spent on investigating new plant products would
only be a small fraction of this aggregate figure.
It is probably to be expected that research in this industry
would follow an extensive-intensive cycle, where new
useful chemical substances are first discovered through
extensive exploration and then developed through intensive
laboratory applications. It is only in the first phase of
pharmaceutical research and development that diversity,
biological and cultural, figures largely; however, from time
to time this input may be crucial for progress to continue.
Finally, it is important to note that the real economic value
lost from possible plant extinctions will be considerably
greater than the financial losses that are identified in these
studies. The market prices do not include the savings to
society in health care and the pain and suffering avoided
through the development of drugs. (In strict terms these
effects should be considered as the marginal difference over
the next best form of treatment.) An estimate of the annual
economic benefits of plant-based drugs currently in use in
the USA gave a range of $34-$300 billion (in 1984 dollars).
This range is very wide because of the wide range in
estimates of the ‘value of a life’ - i.e. actually the value of
a small change in a small risk that affects a very large
number of people. Whatever the precise value, the
economic values involved are clearly very large and are an
order of magnitude greater than the retail market values.
This survey of the value of plant-based pharmaceuticals
demonstrates that there is very real and concrete value
attached to the information derived from genetic variety.
The difficulties in harnessing this value to conserve the
diversity within which it is embedded lie in the
impossibility of knowing which species have the potential
to contribute economic value. Although it is
probabilistically known that these species have substantial
economic value in aggregate, discovering precisely which
species are valuable will take years of extensive research.
There are a number of issues which will have to be
resolved before the market system can develop real
incentives to preserve biodiversity for pharmaceutical
purposes. As it is very difficult to price values as intangible
as information and options, it is necessary to focus on the
creation of mechanisms that can assist in this. These include
the development of patent rights and royalty payments in
natural variety. There is a slowly growing acceptance of the
potential value of biodiversity but there is no real incentive
436
yet to halt the rapid loss of an irreplaceable resource,
despite the economic value that can be attributed to it. The
creation of systems that can recognise and appropriate these
clear but intangible values is a necessary step.
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2. Uses and Values of Biodiversity
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438
Timothy M. Swanson. Authors as follows: Valuing the
environment, Bruce Aylward (LEEC/IIED); Loss of
biodiversity as an economic process, Timothy Swanson;
Current uses of diverse resources, Sarah Gammage;
Ecotourism, Bruce Aylward, Shirra Freedman; Existence
values, Bruce Aylward; Valuation of diverse resource
systems, Sarah Gammage; Preserving future options, Bruce
Aylward, Sarah Gammage; Crop insurance, Timothy
Swanson; Agricultural genetic diversity, Bruce Aylward;
Pharmaceuticals, David Hanrahan.
PART 3
CONSERVATION AND MANAGEMENT OF BIODIVERSITY
The first parts of this book outlined the nature and status of selected elements of
biological diversity (Part 1), and then discussed the uses made of plants and animals and
the economic values which can be associated with biodiversity (Part 2).
Part 3 will introduce some of the policies, systems, institutions and practices employed
in the conservation and management of biodiversity. The eight chapters are grouped into
four principal sections.
The first section treats two principal mechanisms for management: national legislation
(Chapter 28) and protected area systems (Chapter 29). The latter also covers sites which
are components of international protected area systems. These particular topics have been
selected from among the many national-level approaches because of their direct impact
on biodiversity management. Because almost all the elements of which biological
diversity is comprised occur within national boundaries, national policies are self-
evidently central to conservation.
The second section focuses on international policies and instruments which are intended
either to support national approaches, or to deal with resources which lie outside national
boundaries and thus demand international management. Within this section, Chapter 30
includes a tabulation of existing multilateral treaties, and outlines some of the formal
procedures involved in their genesis; many deficiencies exist in the effectiveness of these
treaties but a handful have come to be of considerable global significance. Chapter 31
discusses some ways in which international policy and legal initiatives have supported
national efforts, or could increasingly do so, while Chapter 32 covers additional
assistance which is directly financial in nature (this chapter includes an attempt to
discover to what extent biodiversity is explicitly identified as a sector for aid support).
Finally in this section, Chapter 33 details two examples (fisheries, Antarctica) where
international measures have been designed, with varied success, to manage international
resources.
Section three (Chapter 34) moves to a different viewpoint, away from policy and legal
issues, and focuses on current practices in biodiversity conservation and the institutions
involved in implementing them. Emphasis is placed on threatened species and genetic
resources. In situ and ex situ approaches to plant and animal conservation are compared,
and the need for coordinated planning at national and international levels is stressed.
The fourth and final section (Chapter 35) outlines the origin and development of the
Convention on Biological Diversity. Negotiations to date have been difficult, as
participating countries have a wide variety of perceptions of the role of such a
convention. Some see it purely as a mechanism for ensuring the maintenance of
biodiversity as part of the global heritage, while others regard it as a means of increasing
the returns from genetic resources within their boundaries and ensuring a more equitable
distribution between countries of the costs and benefits derived from maintaining
biodiversity.
he ‘
28. NATIONAL LEGISLATION
Conservation action typically is carried out within policy
and legal systems established by national governments (or
in a few instances, by regional or provincial governments).
With the exception of Antarctica, virtually all the world’s
terrestrial biodiversity occurs within national boundaries and
measures taken by national governments are thus of
fundamental significance.
A wide range of different national policy and legal
measures for the conservation of biodiversity exists which
vary from country to country depending on the social,
political and economic environment. Despite this variety,
there are a number of common legislative techniques in use
throughout the world; this chapter will describe some of the
more important of these. National legislation in this area is
often divided along sectoral lines, with different legislation
covering the protection of flora, fauna and habitats.
THE PROTECTION OF WILD FLORA
The conservation of wild flora has generally had a rather
low priority. As a result, initiatives and legislation at the
national level for the specific protection of wild flora are
rare and on the whole confined to the developed world.
Most European countries have now adopted legislation to
protect wild plants. In the USA, endangered species of wild
flora are protected under the federal Endangered Species
Act and certain States have enacted additional legislation. In
other parts of the world, comprehensive legislation for the
conservation of wild flora exists, for example, in Israel,
Canada, most Australian states and South Africa. Certain
other countries protect wild flora through legislation on
forests. This is commonly the situation in Africa.
Experience has shown that the degree of protection afforded
to wild flora through such legislation is very limited.
Four types of measures common to many countries that
have enacted legislation for the protection of wild flora are
described below.
Collection and possession
The earliest form of legislative protection specifically for
wild flora was restriction on the collection of specimens.
The first such restriction was imposed on the collection of
edelweiss Leontopodium alpinum in the Swiss canton of Zug
in 1911. Most countries which have such legislation have a
differentiated system of protection, with some species being
fully protected and others receiving partial protection. Full
protection of wild flora is normally provided to plants
which have been ‘listed’ under the relevant legislation. The
legislation typically includes prohibitions on taking,
destroying or damaging plants of listed species or any part
of them. Full protection for listed wild plants is, however,
normally limited to public land and exemptions to the
prohibition on collection are usually granted for scientific or
educational purposes.
In some countries, legislation provides for the protection of
all species in certain areas, as opposed to specific plants. In
Austria, for example, collection prohibitions apply to the
441
National Legislation
alpine flora of several mountain regions. In the Swiss
canton of Ticino there is a general prohibition on the
collection of flora in marshes and peatbogs and on river
banks and lake shores. In addition, collection is banned
from certain areas designated because of their scientific
interest. In Italy, collection of all plants growing on rocks
or wetlands in certain areas is banned. In South Africa and
Swaziland there is a complete ban on the collection of wild
flora along public highways for a distance of about 100m on
either side of the road, and several US States have
prohibited the removal of plants along public highways.
Partial protection, in many countries, takes the form of a
ban on mass collection or destruction of wild flora without
good reason. Examples of this type of restriction exist in
Luxembourg, Zimbabwe and parts of Australia. In the UK
there is a general prohibition on uprooting wild flora,
except by landowners, persons authorised by them or by
local authorities. In other jurisdictions (parts of Italy and
Switzerland) there is an additional prohibition on picking
the aerial parts of plants except in limited numbers. In some
areas one is permitted to pick no more than a small bunch
whilst in others the root or bulb of the plant is protected but
gathering of the aerial parts is allowed without limit. Some
jurisdictions (e.g. Belgium, Czechoslovakia and parts of
Austria) which have adopted the latter approach also
stipulate that care must be taken not to damage the root
when picking the flower.
Because of the difficulty of catching offenders in the act of
collecting, the control of possession is a necessary
complement to prohibiting collection and legislation usually
restricts both activities.
One of the common problems with controls on collection is
that they are often limited to public land. On private land,
the owner or occupier may generally collect the flora
growing on that land without restriction and other collectors
need only seek the permission of the owner or occupier.
This is the situation in most common law countries. In the
UK and South Africa, the general restrictions on collecting
and uprooting of all species of wild flora are not applicable
to landowners. In the USA, under the Endangered Species
Act the collection of listed species is only prohibited on
federal land. Wild flora outside federal land is not covered
by this Act, unless the same species are also protected by
State legislation applicable to private land, or are collected
in the course of the violation of a State trespass law; in this
case, under the 1988 amendment to the Endangered Species
Act State offences automatically become federal offences as
well.
The reason that legislative protection of wild flora rarely
extends to private land is because plants are normally
considered the property of the landowner and any attempt
to curtail the use of this property is seen as an infringement
of property rights.
Trade restrictions
Another common legislative mechanism used for protection
of wild flora is legislation imposing restrictions on its trade.
3. Conservation and Management of Biodiversity
The extent of restriction varies considerably from one
country to another. Some national laws contain exhaustive
lists of prohibited activities (e.g. banning possession,
transport, exhibition, offer for sale, sale, purchase); others
merely state that the sale and/or possession of protected
plants without a permit is prohibited. Where the purpose of
legislation is not to prohibit trade altogether but to ensure
the rational utilisation of a natural resource, fairly complex
permit systems have sometimes been developed.
The aim of trade controls is usually to reinforce collection
bans by eliminating the economic incentives for unlawful
taking of wild flora. Thus, many legal systems completely
prohibit trade in fully or partially protected species.
Trade may be prohibited to prevent the exploitation of
certain plants for profit whilst collection for personal use
remains legal. In several Swedish counties, for instance,
certain species may be freely picked but not sold. The
Belgian plant protection order of 1976 contains a list of taxa
in respect of which only collection for commercial purposes
is prohibited. In Costa Rica there is a trade ban on all
species of orchid but no restrictions on collection.
Trade restrictions are usually implemented by requiring
permits for the commercial collection and sale of wild
plants. They are designed to prevent over-exploitation and
to ensure the rational utilisation of economically valuable
plants. In France, for instance, the 1982 Plant Protection
Order contains a list of species which may only be collected
for commercial purposes under a permit from the Ministry
of the Environment. In Italy, the commercial collection and
sale of medicinal plants is also subject to the granting of a
permit. Other examples are found in the legislation of most
Australian states and of Zaire which provides for a licensing
system for the collection of Rauvolfia species. Other
jurisdictions are now attempting to bring under control the
commercial exploitation of a large variety of wild plants
and forest products, such as berries, fungi and mosses,
which until recently were considered almost everywhere as
a free product of nature.
As enforcement is usually difficult, the legislation tends to
be complex. For instance, under the Californian Desert
Native Plants Act of 1981 collection permits are issued by
the local counties. Permits specify the species which may be
harvested, the area from which they may be harvested and
the collection methods authorised. The number of specimens
that can be taken by the permit holder may also be
specified. In addition to the collection permit, the
permission of the landowner must be sought. Detailed
information tags are issued with the permit and must be
attached to the harvested specimens from the time of
collection until they reach their ultimate owner. The owner
must retain the tag as proof of ownership. This elaborate
system is a rather expensive form of conservation which
limits its use to a relatively small number of species and
countries.
Destruction
Many countries have enacted prohibitions on the destruction
of protected or listed species. There are, however, often
serious flaws in this type of protection which limit its
442
effectiveness. The prohibition is often expressed in such
vague terms as to be very difficult to enforce, and is often
limited by so many exceptions that the ban is of little
practical use.
The prohibition also rarely extends to the habitat of wild
flora. One example where controls do extend to include the
habitat of the species is the US Endangered Species Act.
Under this Act federal agencies are not only prohibited
from carrying out any activity which is likely to jeopardise
the existence of listed species, they are also prohibited from
carrying out any action which may result in the destruction
or adverse modification of their critical habitat.
Other examples of specific connections between protection
for a particular species of wild flora and protection of their
habitats exist. In Norway, for example, Article 9 of the
Nature Protection Act of 1970 prohibits development,
construction, pollution and other encroachments in areas of
major importance for protected species to preserve the
habitats. This provision has been applied to Mistletoe
Viscum album, a rare plant in Norway, and an order of
1976 prohibits the felling of trees on which this plant
grows.
Controlling the introduction of exotic species
The introduction of new exotic species can have drastic
consequences for native flora, fauna and natural habitats,
and exotic species pollution is an important threat to
biodiversity in many parts of the world. Preventative action
is essential and legislation controlling deliberate
introductions has now been adopted in many countries. The
system of control is usually regulated by quarantine laws.
Typically these will allow the importation of exotic species
only for limited purposes such as for zoological or botanical
gardens or for research purposes and in many cases only
after it is ascertained that specimens are disease-free.
Commonly, the importation of such species is restricted to
a limited number of entry points in a country where the
customs officials have the capacity to investigate the
consignment to ensure that it complies with the law.
Importation of endangered exotic species is in most
countries subject to additional controls under legislation
implementing the obligations of the Convention on
International Trade in Endangered Species of Wild Fauna
and Flora (CITES) (see Chapter 31).
In certain cases concern over the inadvertent introduction of
exotic species has also caused the promulgation of some
rather drastic measures. In some countries there is a
complete prohibition on the import of certain potentially
harmful species. Many countries have also enacted strict
requirements as to packaging of imports in order to prevent
accidental introduction of invertebrates. The determination
of some countries to keep exotic species pollution to a
minimum is well illustrated by Australia, where even ship
discharges are now regulated in order to prevent the spread
of toxic algal blooms.
THE PROTECTION OF WILD FAUNA
The protection of wild fauna has generally been given much
more attention than the protection of wild flora. Specific
legislation for the protection of wild animals has existed for
many centuries and the legislative mechanisms used are
often very similar to those described above for wild flora.
Indeed, most modern examples of species-specific
legislation cover both flora and fauna.
Taking
The oldest and most common form of protection for wild
fauna has been restriction on taking. Such legislation has
existed in some areas for many hundreds of years
(restrictions associated with medieval hunting reserves in
Europe, for example). In most countries there is a
differentiated system of protection, with some species being
fully protected and others only partially so. Typically, this
differentiated protection is implemented through the use of
appendices containing lists of the species at different levels.
Usually the degree of protection a species receives is
proportional to the seriousness of the perceived threat to its
survival.
Partial protection can vary from strict controls which in
practice are little different from those applied to fully
protected species, to cases where the restrictions have little
practical effect. A typical example of this system can be
found in India, where the principal legislation is The
Wildlife Protection Act 1972. Under this Act there are five
Schedules. Species listed in the first Schedule are fully
protected; those in the other schedules are provided varying
degrees of protection. Species listed in Schedules II, II or
IV are protected from hunting except in accordance with a
licence issued by the relevant government official. The Act
provides for the following kinds of licences: special game
hunting licences for Schedule II species, big game hunting
licences for Schedule III species and wild animal trapping
licences for Schedule IV species. Any Schedule II or III
species which is killed, wounded or captured must be
reported to an authorised government official in accordance
with specified procedures. No licence is required to hunt
Schedule V species. The Act prohibits the hunting of any
wild animals in wildlife sanctuaries and national parks. The
chief warden of such an area may, however, permit hunting
with the prior approval of the state government, provided
it is necessary for the better protection of wildlife in the
particular sanctuary or park.
Legislation for the partial protection of wild fauna is also
commonly found in the controls placed on recreational
hunting and fishing. These laws typically attempt to limit
the taking of species to sustainable levels. Common
techniques employed to achieve these objectives are the
creation of seasons which limit hunting to certain times of
the year, prohibition on taking in certain areas (such as
national parks and game reserves), limitations on the types
of equipment which can be employed, licensing of operators
and establishment of total catch to try to maintain stocks.
Many countries also have elaborate legislative controls for
the commercial exploitation of wild species. Typical of this
is the control exercised over the fishing industry (see
Chapter 33 for some international examples). Here the
controls, although different in degree, are similar to the
types of control exercised over recreational hunting.
443
National Legislation
Possession and trade
Another common form of legislative protection is restriction
on trade of wild species and their products. Frequently
these controls are provided for in the same legislation as
that which controls taking. This type of control usually
operates on a permit basis and these are granted to specific
persons normally on a restricted basis which enables them
to trade in a specified number of animals or their products.
The extent of the restriction varies considerably not only
from country to country but also from species to species
within a particular country. The aim of such restrictions is
the same as in the case of wild flora, i.e. to restrict the
economic incentives for unlawful taking of protected
species. As with restrictions on the trade of wild flora, a
necessary addition to this type of control is restriction on
possession. Thus, most legislation which establishes
restriction on the taking of wild fauna also restricts
possession of such species and their products.
Controls on the import and export of wild fauna also play
an important role in the protection of threatened species by
reinforcing the effectiveness of the trade controls that exist
in a country. Legislative efforts in this regard are
influenced by the work of CITES and in quite a few cases
are confined to implementation of national obligations
arising from CITES. In New Zealand, for example, import
and export of wildlife is principally regulated by the Trade
in Endangered Species Act of 1989 which was specifically
enacted to implement CITES. The Act regulates trade in
endangered, threatened and exploited species identified in
one of the three schedules, which are equivalent to CITES
Appendices I, II and Ill. Any person wishing to trade in
any specimen of such species must apply to the government
for the appropriate permit or authorisation. With regard to
obtaining the necessary permits, separate conditions apply
to export, import, re-export or introduction from the sea of
endangered species, threatened species and exploited
species. In general a permit authorises the holder to
undertake on one occasion the type of trade to which the
authorisation relates. Such permits are non-transferable and
remain in force for six months unless revoked or
surrendered. The Act also provides for extensive powers of
inspection and gives customs officers broad powers of
search and seizure with respect to listed species being
traded in contravention of the Act.
One of the most extensive and innovative regimes
established to control the import and export of wildlife is
found in the USA. The two principle pieces of legislation
establishing this regime are the Endangered Species Act
1973 and the Marine Mammal Protection Act 1972. The
Endangered Species Act 1973 makes it illegal for any
person to import or export endangered species within the
US, to take endangered species within the US or territorial
seas of the US, to take endangered species upon the high
seas, or to sell or offer for sale any endangered species in
interstate or foreign commerce. The Act also makes it
unlawful for any person subject to US jurisdiction to engage
in any trade in specimens or to possess any specimens in
violation of CITES. This Act comprehensively implements
the obligations contained in CITES. These legal norms are
also backed up by extensive administrative resources which
3. Conservation and Management of Biodiversity
ensure the practical implementation of the Act. An unusual
feature of this Act is the extension of its requirements
outside the US itself. Thus US nationals are still bound by
the requirements contain in this Act even though they
themselves may be outside the country.
The Marine Mammal Protection Act 1972 prohibits the
taking on the high seas of marine mammals by any persons
or vessel subject to US jurisdiction; the taking of such
animals by any person in waters or on lands subject to US
jurisdiction; and the importation of marine animals,
products and parts. The Act also has the extraterritorial
application of the Endangered Species Act. One of the main
purposes of the Act is to control commercial exploitation in
order to reduce to insignificance the incidental killing or
serious injury of marine mammals as a result of commercial
fishing operations. To this end, the Act provides that it is
to be administered for the benefit of protected species rather
than the benefit of commercial exploitation. The Act
therefore represents an unusual primacy of conservation
over commercial interests. The Act provides a scheme to
determine the number and kind of marine animals which
can be taken incidentally to commercial fishing, which in
practice essentially requires commercial fishing operations
to adopt modern techniques and equipment to reduce the
hazard to protected species. The products of commercial
fishing operations which are conducted in contravention to
this scheme are banned from importation into the US.
Several such bans have been implemented, the most notable
in relation to control of tuna products because of the
incidental killing of small cetaceans. This type of control of
commercial interests, ensuring that they take account of
protected species, is quite unusual but if properly
implemented a very important means of protection for wild
species.
LIMITATIONS OF SPECIES LEGISLATION
There are a number of common problems with species-
specific legislation. The usual method for providing legal
protection to species consists in laying down prohibitions or
restrictions together with penalties for non-compliance. The
species to which these rules apply are usually listed in an
annex or schedule to the legislation. Normally, the
appropriate government minister, or other authority, is
empowered to amend the list of species by statutory
instrument, thus making it unnecessary to go through the
elaborate process of adopting a new act each time a change
in the list is required. There are usually no criteria laid
down for listing or delisting the species, and so this remains
entirely at the discretion of the particular authority.
An analysis of wildlife protection legislation shows that in
most cases the lists of protected taxa are relatively short,
rarely exceeding 100 entries. Often the lists are largely
dominated by spectacular species attractive to collectors or
the public and do not comprehensively cover the threatened
species in a particular country. The extent of coverage for
plants and invertebrates is frequently very limited. For
instance, a recent survey of plant legislation found that only
five jurisdictions (including France, Greece and Hungary)
protect a large number of species, and often the list reflects
the personal bias of the people working in the relevant
authority. This situation points to the need for public and
accountable procedures for listing.
One example where this is the case is the US Endangered
Species Act. This provides for a detailed listing procedure
involving a preliminary listing, an inquiry and, if requested,
public hearings. An unusual feature of this procedure is that
it can be initiated by any interested person. Only those
species which are determined by the Secretary of the
Interior to be endangered or threatened may be listed. Plant
protection legislation in individual US States usually uses
the same listing criteria but procedural requirements are
generally simpler.
A common problem with much species-specific legislation
is the restricted definition of taking. In some cases the
definition is so narrow as to limit severely the effectiveness
of the legislation. However, perhaps the most important
deficiency with most legislation of this type is the absence
of any provisions for the maintenance of the habitat of the
species. This is despite the fact that protecting critical
habitats is universally recognised as a basic requirement for
species preservation. Even where there is such a provision
it is usually in such general terms that implementation or
enforcement is difficult. Again a notable exception to this
is the US Endangered Species Act.
Species-specific legislation is thus fraught with many
problems and of limited efficacy in the conservation of
biodiversity. Consequently it is only really effective for
species primarily affected by excess exploitation, or as a
last resort measure for rare and endangered species.
THE PROTECTION OF NATURAL HABITATS
The most important form of legislative measure for the
conservation of biodiversity is that for the protection of
ecosystems and habitats. Control over the use of land is the
essential means by which such systems are managed and
protected. National legislation is the most common way for
these controls to be established. There are several common
types of such mechanism.
Protected species habitats
The US Endangered Species Act was cited above as an
example of legislation which extended to protection of
habitats, in this case ‘critical habitats’ of threatened species,
these being defined as areas which are essential to the
conservation of the species concerned. These areas must be
designated and their boundaries precisely described in the
Federal Register. As of October 1987, of the 168 listed
species of wild flora there were 23 species for which
critical habitats had been designated.
The critical habitat concept has also been used in the 1988
Flora and Fauna Guarantee Act of Victoria, Australia.
Under this Act, where a critical habitat designation is made
landowners are prohibited from collecting protected flora in
the critical habitat. The Act also gives the Minister power
to make interim conservation orders prohibiting or
regulating any activity which takes place within or could
have adverse effect on the designated critical habitat. An
order may also contain a positive requirement that specified
works or activities be undertaken. Interim conservation
orders must be complied with by all persons and may be
applicable to any land. However, the designation can only
be made for a period of two years. Before it expires, the
Minister must take all reasonable steps, including the
conclusion of management agreements, to ensure the long-
term conservation of the taxa, communities or critical
habitats for which they were made.
The French Nature Conservation Act of 10 July 1976
contains a general provision prohibiting the destruction,
alteration or degradation of the habitat of protected species.
A decree adopted in 1977 to implement the Act provides
that the central government representatives (préfets) may
make regulations to promote the conservation of the habitat
of listed protected species. The establishment of these
protected areas, known as arrétés de biotope, is not
automatic. Three conditions have to be fulfilled. There must
be an individual order from the préfet designating a certain
area where particular prohibitions apply. The order may
only prohibit activities that can affect the habitat of a
species. It may only apply to a protected species, that is to
say to a species listed in regulations made by the Minister
of the Environment. Subject to these limitations the powers
of the préfet are quite broad, as he may prohibit or
otherwise regulate activities such as vehicle traffic, farming,
drainage, construction or any other action which may be
detrimental to the conservation of the species habitat. No
compensation is provided to landowners.
An important feature of the arrétés de biotope is the
flexibility and simplicity of the procedure underlying their
adoption. In contrast to the establishment of nature
reserves, which requires a long and protracted consultation
procedure, the arrétés de biotope may be adopted with a
minimum of formalities. They are, therefore, increasingly
used as a substitute for nature reserves, which are meeting
with growing opposition from local populations and
authorities.
The network of arrétés de biotope began to develop after
1982-1983. Most of the areas so protected are designed to
preserve the habitat of animal species, for instance
heronries, and the number which exclusively concern plants
is still small. Examples are a few peatlands harbouring rare
and specialised flora species (e.g. Andromeda polifolia,
Drosera spp.) and certain sites of botanical interest
containing species such as Gagea bohemica, Gagea lutea
and Crambe maritima.
Protected areas
The world’s protected area network, the status of which is
examined in Chapter 29, plays a vital and essential role in
the conservation of habitats and ecosystems. With 169
countries in the world having recognised protected area
networks, their use for the conservation of biodiversity is
universal. Whereas the initial purpose of many such areas
was to protect spectacular scenery and provide recreational
facilities, in recent years the concept has evolved to
encompass habitats of endangered species and ecosystems
rich in biodiversity. Even though the legislation used to
establish such areas varies technically from jurisdiction to
jurisdiction the mechanisms used to control or prohibit
certain activities, the essence of the concept of a protected
area, are more or less universal.
445
National Legislation
In countries where there are large tracts of public lands the
establishment of protected areas under public ownership is
relatively straightforward in theory in that the government
can if it so wishes simply manage the area as a protected
area. Unfortunately the simplicity of this solution from a
legal point of view belies the practical difficulties which
often arise. Frequently, the change of management will also
require that control of the land changes from one
government department to another; this change is often
problematic. In some instances it will require legislative
measures to be promulgated, in others cases it will require
the transfer of the property at market prices even though the
‘purchaser’ is another government department.
One simple and effective way to ensure that government
departments preserve natural habitats on public land is the
‘wilderness area’ concept as used in the USA. Pursuant to
the Wilderness Act of 1964 it is possible to ban the
construction of all roads and tracks and other means of
access within a specified area. The National Wilderness
Preservation System, which is made up of these specified
areas, has developed rapidly and is intended to cover some
400,000km?* of federal land under the control of various
government departments. The potential of this type of
measure is obvious because threats typically escalate
following increasing access to wilderness areas by road
construction.
If the land requiring protection is in private hands,
governments have used a variety of mechanisms to establish
the necessary protection. In some instances they have
simply acquired the land from the owner. This mechanism
can be expensive. One way that governments have sought
to ameliorate this cost is to acquire a lessor interest in the
land, such as the right of drainage, where such rights are
separable.
Alternatively governments can and have used their rights of
expropriation to force private owners to either relinquish
the land or agree to controls over the use of the land.
Governments are now reluctant to use such powers
especially for conservation purposes. More commonly
governments will impose restraints on the use of land by
private persons by, for instance, banning all forms of use
which are detrimental to the ecosystems present in the area.
Such forms of control are not always constitutionally
possible, as in common law countries where such a
curtailment of rights is generally perceived as unlawful. In
these countries, the government is generally only able to
impose such controls under a voluntary management
agreement with the owner. Under voluntary agreements the
owner commits himself not to use the land for certain
purposes. One example of this type of agreement is that
found in England where, under the Wildlife and
Countryside Act 1981, English Nature (formerly the NCC)
can enter into agreement with the owners of Sites of Special
Scientific Interest (SSSIs).
Protection of private land is also facilitated by the legal
system through the use of caveats. These rights attach to the
land itself and will bind future owners. Such rights exist in
most common law jurisdictions. In some countries the
3. Conservation and Management of Biodiversity
government has pre-emptive rights over the sale of certain
land should it happen to be sold by the owner. Such
provisions exist in several European countries. In the USA
the government frequently negotiates a pre-emptive right
individually with the owner. In France the pre-emptive right
is also linked with a mechanism to finance the purchase of
such properties which come onto the market. Under the
legislation creating the pre-emptive right the particular
department is also empowered to collect a tax on the
construction of buildings the proceeds of which are
hypothecated to the acquisition of private land.
Land-use controls
Many countries have legislation limiting the use to which
land may be put. Such land-use controls or zoning
restrictions typically control activities such as construction
or mining and are normally restricted to the urban
environment. In a few countries zoning restrictions also
extend to rural areas; however, agricultural and forestry
activities are normally exempted from their provisions.
In a few countries such mechanisms are used to protect
natural habitats. Examples of such mechanisms include:
special protection orders for specific sites; the use of
specially protected areas in local zoning plans; or the
prohibition on altering of certain habitats without a permit.
A prime example of such a mechanism being used to
protect natural habitats is the Danish Nature Conservation
Act of 1969 (as amended). This Act establishes a strict
system of permits applicable to all activities which may
have an adverse effect on river beds, lakes, peatbogs, salt
marshes, coastal vegetation and natural grasslands. This
type of approach has also been adopted in many European
countries, North America and parts of Australia.
Another important and common land-use control is
restriction on felling of private forests. In most cases, the
Testrictions are not applicable to the government forestry
department itself. One exception to this is found in the USA
where, under the US Federal Forests and Rangelands
Renewable Resources Planning Act of 1976 (as amended),
the discretionary authority of the Forest Service is curtailed
and the objective of the organisation must now include the
maintenance of all plants and animal species and the
promotion of the recovery of endangered species. Forest
plans must be drawn up for each unit in the National Forest
System using an interdisciplinary approach and including
public participation. A common problem with this
mechanism is that the purpose of the legislation is often not
the preservation of natural forests but simply the
maintenance of forest cover. This means that the
replacement of native forest, rich in biodiversity, with
446
comparatively sterile monocultures of production timber is
not regulated by such controls.
Incentives
A common legislative mechanism to help conserve natural
habitats is the provision of incentives or disincentives to
influence the activities of land users to conserve natural
habitats. Examples of such mechanisms are the EC
regulation providing for the subsidy payments to farmers to
maintain the natural environment on their land, and the
granting of land tax credits for the preservation of wetlands
or natural prairie areas, or for the conservation of river
banks, in the US State of Minnesota. Another important
example of an incentive, although an indirect one, is the tax
exemptions granted in many countries to many conservation
organisations on the basis of their charitable status. In the
USA, land owned by conservation organisations or land
dedicated to conservation is frequently exempt from land
tax.
Many countries not only provide incentives to preserve
natural habitats but also penalise environmentally harmful
activities. Measures of this sort include the refusal of
subsidies and the imposition of special taxes on such
activities. The UK Wildlife and Countryside Act 1981
contains such a mechanism. It provides that agricultural
subsidies may be refused for activities which will adversely
affect the flora, fauna and physiogeographical features of
national parks or in areas specially designated for that
purpose (e.g. SSSIs). The US Food Security Act of 1985 is
also another example of such a mechanism. The purpose of
this Act is to remove up to 40 million acres (16 million
hectares) of erodible land from agricultural production to,
inter alia, reduce erosion and enhance wildlife. It seeks to
achieve this by removing a number of subsidies from crops
produced on highly erodible soil or altered wetland.
Indirect legislation
The types of legislative mechanisms described above are all
examples of direction protection of biodiversity. In many
countries there exist numerous legislative mechanisms
which while not directly protecting biodiversity do
nonetheless play a vital role in its conservation. Examples
of this type of legislation are pollution control laws or
legislation regulating development and investment in a
country. Such controls can and do have an important effect
on the conservation of biodiversity in a country. If properly
framed, they can be powerful forces for the conservation of
biodiversity; if not, such regimes can have drastic
consequences for its conservation.
Chapter contributed by Sam Johnston.
29. PROTECTED AREAS
Natural ecosystems and the habitats they contain are subject
to some degree of control and protection in every country
in the world. Many different legal and administrative
mechanisms are used by governments to manage habitats
for the conservation of biodiversity. Protected area systems
are central to such management. This section will provide
information on protected areas which contribute to such
systems, charting the growth in protected areas over the
past century. It will also examine the extent to which
different geographic and biogeographic regions, and biome
types are covered by protected area systems, and highlight
major gaps in the network.
NATIONAL PROTECTED AREA SYSTEMS
There is considerable variation between countries in the
mechanisms used to create and maintain systems of
protected areas. Some standard means of classification
needs to be used in making international comparisons. The
IUCN, through its Commission on National Parks and
Protected Areas (CNPPA), has developed a system of
classification for different types of protected area, based
upon management objectives. This system has 10 different
classes of protected areas, two of these, World Heritage
Sites (X) and Biosphere Reserves (IX) being international
designations.
In the analysis in this chapter the term ‘protected area’ is
defined as an area of 1,000ha or more in IUCN
Management Categories I-V, managed by the highest
competent authority. These are the criteria used in
compiling the 1990 United Nations List of National Parks
and Protected Areas (IUCN, 1990).
However, statistics prepared using such standard criteria
omit a range of significant sites. For instance, the statistics
presented here do not include: sites which are in other
management categories (such as multiple-use areas (VIID),
areas under 1,000ha (such as the numerous small reserves
in Europe), areas outside the IUCN Categories altogether,
such as partially protected areas (e.g. hunting reserves), and
areas not managed by the ‘highest competent authority’ but
protected by private organisations (such as NGOs),
superstition, isolation or military activity. All of these
conserve significant amounts of biodiversity. Whilst
information on such sites is available, it is not yet consistent
and has.
The wise management of areas which are devoted to
agriculture, through management techniques such as non
site-specific legal instruments, planning control, voluntary
agreements, and integrating conservation principles into
land-use planning, also play an essential role in
conservation of biodiversity. Indeed, in most countries,
management of land-use outside the national network of
protected areas will play as important a role in the
conservation of biodiversity as will the network itself. In
order to examine comprehensively the role that land
management plays in the conservation of biodiversity, it
would be necessary to survey the use of these other areas
and techniques as well.
447
Protected Areas
Unfortunately this is not possible at this stage because of
the paucity of reliable data on these important measures.
Categories and management objectives of protected areas
The following categories and criteria for protected areas are
abridged from IUCN (1984).
I Scientific Reserve/Strict Nature Reserve: to protect
nature and maintain natural processes in an
undisturbed state in order to have ecologically
representative examples of the natural environment
available for scientific study, environmental
monitoring, education, and for the maintenance of
genetic resources in a dynamic and evolutionary
state.
National Park: to protect natural and scenic areas of
national or international significance for scientific,
educational and recreational use.
Natural Monument/Natural Landmark: to protect
and preserve nationally significant natural features
because of their special interest or unique
characteristics.
Managed Nature Reserve/Wildlife Sanctuary: to
assure the natural conditions necessary to protect
nationally significant species, groups of species,
biotic communities, or physical features of the
environment where these require specific human
manipulation for their perpetuation.
Protected Landscape or Seascape: to maintain
nationally significant natural landscapes which are
characteristic of the harmonious interaction of man
and land while providing opportunities for public
enjoyment through recreation and tourism within
the normal life style and economic activity of these
areas.
Other categories defined by IUCN but not analysed here are
Category VI (Resource Reserve), Category VII (Natural
Biotic Area/Anthropological Reserve) and Category VIII
(Multiple-Use Management Area/Managed Resource Area).
The classes and their different management objectives are
given in Table 29.1.
Development
Areas that are in some sense ‘protected’, in that access or
forms of use are controlled, have existed for many
thousands of years. In India, protected areas have existed
since the 4th century BC, with the establishment of
Abhayaranxyas or forest reserves. In the Pacific region, the
imposition of tapu (taboo) effectively created protected
areas; the existing protected area on Niue, for example,
consists of a tapu forest. Hunting reserves have existed in
Europe for hundreds of years. The first modern examples
of protected areas were established towards the end of the
19th century.
3. Conservation and Management of Biodiversity
Table 29.1 Protected area objectives
PROTECTED AREA DESIGNATION (IUCN CATEGORY NUMBER)
Scientific National Natural Managed Protected Resource Natural Multiple-
Reserve Park Monument Nature Landscape Reserve Biotic Use Area
| i] Wl Reserve Vv Vi Reserve vill
CONSERVATION OBJECTIVE IV vil
Maintain sample ecosystem in e e ° e ° fe) e
natural state
Maintain ecological diversity and ° e e ° ° fo) e °
environmental regulation
Conserve genetic resources e ° ° e °
Provide education, research and ° ° ° ° °
environmental monitoring
Conserve watershed, flood control ° e ° [o) fo) ° ° °
Control erosion and sedimentation ° fo) ° fo) oO ° fe)
Maintain indigenous use or e fo) (0)
habitation
Produce protein from wildlife fo) fo) fo) °
Produce timber, forage or ° fo} ° e
extractive commodities
Provide recreation and tourism e e ° e ° e
service
Protect sites and objects of ° fe) ° e ° e °
cultural, historical, or
archaeological heritage
Protect scenic beauty ° e ° ° e
Maintain open options, ° e
management flexibility,
multiple-use
Contribute to rural development ° e ° ° e ° fo) e
Sources: Miller, K.R. 1980. Planning National Parks for Ecodevelopment, Center for Strategic Wildland Management Studies, Ann Arbor;
TUCN/UNEP 1986. Managing Protected Areas in the Tropics. TUCN, Gland, Switzerland.
Notes: © = Primary Objectives, o = Compatible Objectives.
By the beginning of this century many countries had either
already established protected areas or were contemplating
doing so. The concept, however, was slow to develop to a
stage where any one country had developed a
comprehensive network of actively managed protected
areas. It was not until the 1940s that protected areas were
beginning to be established in any significant number. After
World War Il, the number of protected areas established
continued to be low, and the rate at which land was being
incorporated into the system did not increase above pre-
World War II levels until the early 1960s. In 1962 the
establishment of protected areas began to increase
dramatically. An important stimulus for this increase may
have been the first World Parks Congress held in Seattle,
USA in 1962. This meeting signified the emergence of the
modern protected area network with over 80% of the
world’s protected areas being established since then. Table
29.2 shows that the increase experienced during this period
has continued unabated until the present day.
The rates of growth of protected areas on global and
regional bases are illustrated in Fig. 29.1 and 29.2
respectively, showing the number of sites and the area
protected. It should be noted that the creation of Greenland
National Park in 1974, which covers some 97 million ha,
and the creation of Great Barrier Reef Marine
448
Table 29.2. Dates of establishment of
protected areas
DATES NUMBER AREA (km?)
Pre-1962 1,433 1,324,600
1962-1971 1,372 862,800
1972-1981 2,258 3,559,800
1982 onwards 2,140 1,684,100
Date unclear 1,288 303,600
TOTAL 8,491 7,734,900
Park in the 1980s, which covers some 34 million ha, has a
marked effect on the area protected for the relevant period.
Fig. 29.2 shows that there is considerable regional variation
in the development of protected area networks. Reasons for
this variation include: cultural and historical factors, the
development of interest in wildlife and conservation in the
region, and patterns of settlement and land-use.
Despite the regional variations the graphs do illustrate a
number of global trends. For most regions, networks of
protected areas are a recent phenomenon, with only Africa
Protected Areas
Figure 29.1 World growth of the protected areas network
2,000
1,500 HEIGHT + Greenland National Park
Area (x1000 km?) * Great Barrier Reef Marine Park
1,000
1875 1890 1900 1910 1920 1930 1940 1950 1960 1970 1980 19390
1880 1895 1905 1915 1925 1935 1945 1955 1965 1975 1985
Five year period ending.....
ques §=6Number of sites
Area (x1000 km?)
1875 1890 1900 1910 1920 1930 1940 1950 1960 1970 1980 1930
1880 1895 1905 1915 1925 1935 1945 1955 1965 1975 1985
Cumulative growth to.....
449
3. Conservation and Management of Biodiversity
Figure 29.2 Regional growth of the protected areas network
===—<—= Number of sites
Area (x1000 km?)
North
and
Central America
* Greenland National Park
USSR (former)
South America Africa
300 300
Oceania
200 200 * Great Barrier Reef Marine Park
100 100
fe) ES ES BS B38 iC fo} eee ef) i Ek.
g AQF Qt 1929 493° QAO .96 1969 191 1980 ,990 as 49 49° 1929 193 .qA% .969 ,96° 1919 .989 .990
Five year period ending..... Five year period ending.....
450
and North America having a significant history of protected
areas before 1962. The rates of growth for both number of
sites and area protected are still high throughout much of
the world. There is a tendency for larger protected areas to
be established in the developing world.
Present status of national systems
Protected areas meeting the criteria given now exist in 169
countries in the world. There are currently some 8,491 sites
covering some 7,734,900km? or some 5.19% of the earth’s
land area. The largest protected area is Greenland National
Park, which covers 972,000km?. In 115 countries, 1,328
sites covering some 3,061,300km? have marine or coastal
elements within them. Of these, 94 sites have coral reefs.
The largest marine protected area is Great Barrier Reef
Marine Park, which covers some 340,000km?.
The relative proportion of each type of IUCN Category is
illustrated in Fig. 29.3. Managed nature reserves/wildlife
sanctuaries (Category IV) are the most prevalent type of
protected area in terms of number of sites. National parks
(Category II) cover more area than any other category. The
extent to which each category is applied varies considerably
from region to region, as a result of cultural, demographic
and geographic factors. Thus, in Europe, where there is
very little natural habitat and where man has extensively
altered that which remains, most large protected areas are
managed as protected landscapes. In Australia, where man’s
influence is less pervasive, the predominant protected area
category is the national park.
The size distribution of protected areas is illustrated in Fig.
29.4. The most common size for a protected area on a
worldwide basis is only 10-30km?. However, the majority
of the world’s 7.7 million km? of protected area is
contained in a relatively few large sites. These figures
suggest that fragmentation may be a problem in providing
protection to many of the world’s natural habitats.
Significant regional differences in size distribution can also
be deduced from Fig. 29.2.
Table 29.3 presents the distribution of protected areas
according to the World Bank classification of the country’s
economy. The classes are based on per capita income. The
low income class is subdivided according to country size
Figure 29.3 Protected areas by IUCN
category
Area covered
Number of sites
451
Protected Areas
(‘large’ includes India and China). The ‘middle income
(upper)’ class is distorted by the former USSR, where the
protected areas network has extremely low coverage and the
country area is very large. The high income class is divided
by membership in the Organization for Economic
Cooperation and Development (OECD). The extraordinary
figures for the Non-OECD group are because of Greenland
National Park. Protected areas are fairly evenly spread by
income groups and quite high even for very poor countries.
The smaller average size of a protected areas for the large
low income group and the OECD countries probably
reflects the high population densities of these countries.
A major objective of the protected area system of the world
is maintaining the diversity of species and ecosystems.
Biogeographical analysis of protected area coverage
provides information on how effectively the various natural
ecosystems are being conserved.
A basic system of biogeographic analysis has been worked
out for terrestrial ecosystems by Udvardy (1975). He
divides the world into eight biogeographical realms,
continent or subcontinent-sized areas, which are further
divided into 193 provinces defined by significant differences
in flora, fauna, or vegetation structure. The provinces are
associated with 14 biomes, which are major regional
ecological communities of plants and animals. It should be
noted that a protected area located within a particular
province may not necessarily contain vegetation typical of
that province. Thus a protected area within the Congo
Rainforest province may not necessarily contain tropical
humid forest, and although insular Malaysia, Indonesia and
the Philippines are classified as mixed island systems, they
all contain extensive tropical humid forests.
Table 29.4 presents the extent to which each biome is
covered by protected areas. This table shows that temperate
grasslands and lake systems are poorly represented in the
protected area network, and that this is an area requiring
attention. The conclusions that can be drawn from the high
level of aggregation at the biome level are limited. A more
accurate picture of ecosystem protection can be gained from
an analysis of protected area coverage at the province level.
Table 29.5 lists in descending order the percentage
coverage of each province. The analysis of protected area
coverage at this level still suffers from the problems
mentioned above, albeit in a reduced way. These data are
also presented in map form in Fig. 29.5.
Fig. 29.6 illustrates the distribution of marine and coastal
protected areas throughout the world. There are also 559
sites that have an altitudinal range of 1,500m or more and
Fig. 29.7 illustrates the distribution of these mountainous
areas.
Studies of protected area coverage at regional and national
levels would provide a much better assessment of priorities,
and many such studies have been undertaken. The
mechanisms for assessment used in these studies vary very
widely, so an assessment of coverage based on these studies
has not been attempted. A range of regional studies have
been published by IUCN and others.
3. Conservation and Management of Biodiversity
Figure 29.4 Protected areas by size class frequency
3,000
2,500
2,000
1,500
1,000
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Table 29.3 Distribution of protected areas by World Bank income groups
Number % of Area % of Average Country % of
Total (km?) Total Size Area country
INCOME GROUP AND No. Area (km?) (km?) Area
SUBGROUP
Low Income (large) 758 8.9 421,300 5.5 556 12,764,000 3.3
Low Income (small) 734 8.6 1,067,300 13.8 1,454 24,636,000 4.3
Middle Income (lower) 1,051 12.4 1,338,500 17.3 1,274 23,173,000 5.8
Middle Income (upper) 1,126 13.3 1,200,400 {EBS 1,066 41,404,000 eed
High Income (OECD) 4,713 55.5 2,677,100 34.6 568 31,079,000 8.6
High Income (Non-OECD) 62 0.7 990,600 12.8 15,977 2,381,000 41.6
Income not assigned 47 0.6 39,700 0.5 845 13,677,000 0.3
TOTAL 8,491 100.0 7,734,900 100.0 911 149,114,000 100.0
Table 29.4 Distribution and coverage of protected areas by biome type
BIOME TYPE PROTECTED AREAS BIOME % OF
NUMBER AREA AREA TOTAL
(km2) (km?) AREA
Subtropical/temperate rainforests/woodlands 935 366,100 3,928,000 9.32
Mixed mountain systems 1,265 819,600 10,633,000 7.71
Mixed island systems 501 246,300 3,244,000 7.59) **
Tundra communities 81 1,643,400 22,017,000 7.46
Tropical humid forests 501 522,000 10,513,000 4.96
Tropical dry forests/woodlands 807 818,300 17,313,000 4.73
Evergreen sclerophyllous forests 786 177,400 3,757,000 4.72
Tropical grasslands/savannas 56 198,200 4,265,000 4.65
Warm deserts/semi— deserts 296 957,700 24,280,000 3.94
Cold—winter deserts 139 364,700 9,250,000 3.94
Temperate broad — leaf forests 1,509 357,000 11,249,000 3.17
Temperate needle—leaf forests/woodlands 440 487,000 17,026,000 2.86
Lake systems 18 6,600 518,000 1.28
Temperate grasslands 196 70,000 8,977,000 0.78
Classification unknown 961 700,800 0 NA
TOTAL 8,491 7,734,900 146,968,000 5.26
Notes: ** Protected area includes significant marine areas, inflating the % figure. Biome definitions after Udvardy, 1975.
452
Protected Areas
Table 29.5 Distribution and coverage of protected areas by biogeographic
province
PROVINCE REALM PROTECTED AREA PROVINCE % OF
NUMBER AREA AREA TOTAL
(km2) (km2) AREA
Cocos Island Neotropical 1 24 24 100.0 **
Everglades Neotropical 17 8,080 6,800 100.0 **
Fernando De Noronja Island Neotropical 1 362 17 100.0 **
Galapagos Islands Neotropical 1 7,600 7,600 100.0 **
Aleutian Islands Nearctic 9 79,100 124,500 63.5 **
Campos Limpos Neotropical 6 108,640 207,300 52.4
Sitkan Nearctic 85 172,010 350,500 49.1
Arctic Desert and Icecap Nearctic 2 982,500 2,119,500 46.4
Valdivian Forest Neotropical V2) 40,160 111,900 35.9
Lesser Antillean Neotropical 26 2,240 6,600 34.0 **
Alaskan Tundra Nearctic 25 309,420 958,500 32.3
Chilean Nothofagus Neotropical if 39,200 123,700 31.7
Colombian Montane Neotropical 23 42,500 154,800 27.5
Queensland Coastal Australian 78 81,690 300,200 27.2
Namib Afrotropical 7 95,970 364,600 26.3
Panamanian Neotropical 9 10,350 40,100 25.8
Cocos — Keeling and Christmas Islands Indomalayan 1 87 337 25.8
Ceylonese Monsoon Forest Indomalayan 42 7,760 34,900 22.2
Tasmanian Australian 28 13,910 68,000 20.5
Yukon Taiga Nearctic 27 203,110 1,019,600 19.9
Sonoran Nearctic 38 100,540 507,800 19.8
Kalahari Afrotropical 10 97,770 504,900 19.4
Venezuelan Dry Forest Neotropical 41 50,640 270,300 18.7
Comores Islands and Aldabra Afrotropical 1 350 1,920 18.2
Arctic Desert Palaearctic 5 34,910 195,900 17.8
Seychelles and Amirantes Islands Indomalayan 3 36 204 17.5
Central European Highlands Palaearctic 381 62,620 369,900 16.9
Hawaiian Oceanian 19 2,790 16,700 16.7
Insulantarctica Antarctic 11 3,140 19,200 16.3
Southern Andean Neotropical 51 106,940 662,900 16.1
Ryukyu Islands Palaearctic 3) 391 2,500 15.8
Southeastern Polynesian Oceanian 12 640 4,200 15.4
Venezuelan Deciduous Forest Neotropical 16 8,960 58,900 15.2
Scottish Highlands Palaearctic 43 7,110 46,800 15.2
New Caledonian Oceanian 16 2,510 16,700 15.0
British Islands Palaearctic 103 39,560 266,600 14.8
Serro Do Mar Neotropical 56 35,980 243,800 14.8
Northern Andean Neotropical 18 37,380 256,500 14.6
Cape Sclerophyll Afrotropical 51 18,670 129,700 14.4
Sulawesi (Celebes) Indomalayan 38 25,230 196,700 12.8
Andaman and Nicobar Islands Indomalayan 42 797 6,200 12.8
Sierra—Cascade Nearctic 82 28,380 228,700 12.4
Guinean Highlands Afrotropical 3 9,720 80,000 124
Canadian Tundra Nearctic 20 198,210 1,733,400 11.4
Central African Highlands Afrotropical 7 30,630 269,500 11.4
Neozealandia Antarctic 150 29,050 266,000 10.9
Japanese Evergreen Forest Palaearctic 481 28,370 266,900 10.6
Sumatra Indomalayan 38 49,080 461,900 10.6
Bahamas —Bermudean Neotropical 6 1,350 12,800 10.6
Greater Antillean Neotropical 35 9,990 95,800 10.4
Himalayan Highlands Palaearctic 79 82,570 860,100 9.6
Pamir—Tian—Shan Highlands Palaearctic 30 60,910 643,200 9.5
East African Woodland/Savanna Afrotropical 71 142,820 1,510,600 9.5
Yungas Neotropical 18 44,100 483,100 9.1
Anatolian—Iranian Desert Palaearctic 47 200,300 2,203,800 9.1
Icelandian Palaearctic 23 9,170 101,600 9.0
Iberian Highlands Palaearctic 114 28,450 316,100 9.0
Central Polynesian Oceanian 4 362 4,200 8.7
Rocky Mountains Nearctic 145 127,850 1,578,500 8.1
Macaronesian Islands Palaearctic 11 1,130 14,000 8.0
Taiwan Indomalayan 5 2,890 36,600 7.9
Oregonian Nearctic 32 9,430 124,600 7.6
Amazonian Neotropical 34 184,090 2,509,400 7.3
Malayan Rainforest Indomalayan 23 12,740 179,200 7.1
Papuan Oceanian 36 68,180 960,100 7.1
Java Indomalayan 43 9,730 137,900 7A
Southern Mulga/Saltbush Australian 14 58,700 837,000 7.0
Southern Sclerophyll Australian 67 17,080 246,700 6.9
Western Sclerophy/l Australian 125 26,600 410,800 6.5
Indochinese Rainforest Indomalayan 60 29,090 452,500 6.4
453
3. Conservation and Management of Biodiversity
Table 29.5 Distribution and coverage of protected areas by biogeographic
province (continued)
PROVINCE REALM PROTECTED AREA PROVINCE % OF
NUMBER AREA AREA TOTAL
(km2) (km2) AREA
Thar Desert Indomalayan 39 45,680 711,800 6.4
Cuban Neotropical 34 6,910 109,800 6.3
Lesser Sunda Islands Indomalayan 20 5,390 86,600 6.2
Atlantic Palaearctic 120 44,300 715,900 6.2
Miombo Woodland/Savanna Afrotropical 38 148,400 2,432,100 6.1
Borneo Indomalayan 66 42,310 741,000 5.7
Eastern Sclerophyll Australian 169 35,920 643,800 5.6
Central Desert Australian 17 98,960 1,777,100 5.6
Takla—Makan-— Gobi Desert Palaearctic 19 120,000 2,184,600 5.5
West African Woodland/Savanna Afrotropical 80 177,370 3,247,600 5.5
Caucaso—lIranian Highlands Palaearctic 66 49,940 936,000 5.3
Puna Neotropical 19 23,390 464,900 5.0
Mahanadian Indomalayan 29 10,970 219,400 5.0
Lake Titicaca Neotropical 1 362 7,200 5.0
Colombian Coastal Neotropical 9 11,280 237,200 48
South African Woodland/Savanna Afrotropical 104 80,350 1,694,800 47
Middle European Forest Palaearctic 401 68,870 1,467,300 47
Malabar Rainforest Indomalayan 43 10,140 223,600 45
Northern Savanna Australian 10 26,100 580,900 45
Central American Neotropical 58 13,900 310,000 45
Balkan Highlands Palaearctic 103 9,830 221,200 4.4
Campechean Neotropical 19 11,400 259,200 4.4
Szechwan Highlands Palaearctic 52 24,660 578,600 4.3
Eastern Sahel Afrotropical 4 48,460 1,169,700 44
East African Highlands Afrotropical 11 2,680 65,500 41
Indus— Ganges Monsoon Forest Indomalayan 129 55,680 1,412,200 3.9
Kamchatkan Palaearctic 1 10,990 283,300 3.9
Congo Rain Forest Afrotropical 24 71,900 1,921,900 3.7
Northern Coastal Australian 14 12,890 350,400 3.7
Equadorian Dry Forest Neotropical 4 1,840 50,300 3.7
Canadian Taiga Nearctic 286 180,310 5,127,200 3.5
Pannonian Palaearctic 33 3,520 102,500 3.4
Sahara Palaearctic 17 226,970 6,960,900 3.3
Patagonian Neotropical 25 13,200 413,100 3.2
Ethiopian Highlands Afrotropical 7 16,060 505,400 3.2
Babacu Neotropical 6 9,030 293,000 3.1
Madrean—Cordilleran Nearctic 83 23,410 763,200 3.1
Mediterranean Sclerophyll Palaearctic 227 36,590 1,194,700 3.1
Western Mulga Australian 15 22,610 778,100 2.9
Congo Woodland/Savanna Afrotropical 6 37,740 1,356,800 2.8
Yucatecan Neotropical 3 1,070 40,000 27
Manchu-— Japanese Mixed Forest Palaearctic 167 32,730 1,252,300 2.6
Uanos Neotropical 3 11,410 438,000 2.6
Higharctic Tundra Palaearctic 2 22,290 859,900 2.6
Bengalian Rainforest Indomalayan 20 4,630 179,900 2.6
Chilean Sclerophyll Neotropical 8 1,470 57,300 2.6
Thailandian Monsoon Forest Indomalayan 62 24,530 959,700 2.6
Uruguayan Pampas Neotropical 12 12,890 522,200 25
Guinean Rain Forest Afrotropical 23 14,960 607,000 2.5
Guyanan Neotropical 27 24,860 1,009,100 2.5
Iranian Desert Palaearctic 9 9,810 403,500 2.4
Oriental Deciduous Forest Palaearctic 200 64,100 2,751,400 2.3
Lake Ladoga Palaearctic 1 410 17,600 2.3
Malagasy Rain Forest Afrotropical 16 4,560 200,600 23
Eastern Forest Nearctic 192 49,950 2,223,000 2.2
Mascarene Islands Afrotropical 5 100 4,500 2.2
Chinese Subtropical Forest Palaearctic 78 18,790 863,000 2.2
South African Highlands Afrotropical 38 4,330 199,000 2.2
Altai Highlands Palaearctic 7 22,820 1,048,300 2.2
Austroriparian Nearctic 98 12,200 596,900 2.0
Campos Cerrados Neotropical 25 36,280 1,778,600 2.0
Great Lakes Nearctic 13 5,140 254,500 2.0
Somalian Afrotropical 27 43,280 2,166,800 2.0
Arabian Desert Palaearctic 31 59,300 2,996,100 2.0
Philippines Indomalayan 27 5,730 292,200 2.0
Burma Monsoon Forest Indomalayan 29 5,800 297,200 2.0
Subarctic Birchwoods Palaearctic 13 2,530 132,500 1.9
Eastern Grasslands and Savannas Australian 51 10,080 527,800 1.9
Malagasy Woodland/Savanna Afrotropical 19 6,150 324,100 1.9
South Chinese Rainforest Indomalayan 53 3,540 189,000 1.9
Coromandel Indomalayan 4 1,570 88,400 1.8
454
Protected Areas
Table 29.5 Distribution and coverage of protected areas by biogeographic
province (continued)
PROVINCE REALM PROTECTED AREA PROVINCE % OF
NUMBER AREA AREA TOTAL
(km?) (km?) AREA
Brigalow Australian 12 3,940 231,600 1.7
Californian Nearctic 13 8,650 526,500 1.6
Sinaloan Neotropical 5 2,970 192,100 1.5
Boreonemoral Palaearctic 152 18,460 1,285,300 1.4
Western Sahel Afrotropical 9 39,510 2,814,700 1.4
Chilean Araucaria Forest Neotropical 2 454 32,900 1.4
Gran Chaco Neotropical 14 12,830 988,500 1.3
Karroo Afrotropical 18 4,660 377,700 1.2
West Eurasian Taiga Palaearctic 117 65,400 5,342,600 1.2
Great Basin Nearctic 21 7,230 660,400 11
Micronesian Oceanian 4 23 2,200 14
Chihuahuan Nearctic 19 5,820 577,200 1.0
Brazilian Rain Forest Neotropical 59 15,030 1,533,800 1.0
Mongolian—Manchurian Steppe Palaearctic 17 22,530 2,605,100 0.9
Hindu Kush Highlands Palaearctic 5 1,830 217,100 0.8
East Siberian Taiga Palaearctic 11 45,750 5,536,100 0.8
Guerreran Neotropical 6 1,290 158,400 0.8
Northern Grasslands Australian 6 6,700 967,000 0.7
Monte Neotropical 26 8,490 1,234,800 0.7
Argentinian Pampas Neotropical 17 3,470 512,200 0.7
Turanian Palaearctic 15 13,940 2,116,800 0.7
Lake Ukerewe (Victoria) Afrotropical 1 460 69,500 0.7
Pontian Steppe Palaearctic 25 12,550 1,945,400 0.6
Malagasy Thorn Forest Afrotropical 2 450 70,700 0.6
Madeiran Neotropical 4 10,460 1,671,800 0.6
Lowarctic Tundra Palaearctic 1 13,490 2,158,100 0.6
Deccan Thorn Forest Indomalayan 9 1,940 338,400 0.6
Grasslands Nearctic 69 7,690 2,442,300 0.3
Lake Malawi (Nyasa) Afrotropical 1 87 28,900 0.3
West Anatolian Palaearctic 4 107 37,600 0.3
Aral Sea Palaearctic 1 183 67,500 0.3
Caatinga Neotropical 6 2,430 899,700 0.3
Ceylonese Rainforest Indomalayan 1 80 31,100 0.2
Tamaulipan Nearctic 3 500 210,400 0.2
Atlas Steppe Palaearctic 6 910 421,500 0.2
East Melanesian Oceanian 2 50 27,000 0.2
Pacific Desert Neotropical 2 490 290,400 0.2
Burman Rainforest Indomalayan 2 200 257,600 0.1
Brazilian Planalto Neotropical 3 140 219,200 0.1
Tibetan Palaearctic 3 240 1,268,100 0.0
Maudlandia Antarctic 5 340 10,465,200 0.0
Marielandia Antarctic 1 3 2,194,000 0.0
Arctic Archipelago Nearctic 0 0 690,000 0.0
Ascension and St Helena Islands Afrotropical 0 (e) 200 0.0
Greenland Tundra Nearctic ie) (e) 498,600 0.0
Laccadives Islands Indomalayan 0 0 32 0.0
Lake Baikal Palaearctic 0 0 32,300 0.0 *
Lake Rudolf Afrotropical 0 0 7,300 0.0 *
Lake Tanganyika Afrotropical 0 0 32,800 0.0 *
Maldives and Chagos Islands Indomalayan_ 0 0 36 0.0
Revilla Gigedo Island Neotropical fe) (0) 200 0.0
South Trinidade Island Neotropical 0 (0) 11 0.0
Classification unknown 951 692,830 0 NA
TOTAL 8,491 7,734,900 146,970,700 5.3
Source: WCMC.
Note: * Shorelines are often protected but not included in this figures. ** Protected area includes significant marine areas, inflating the
% figures. Province definitions after Udvardy, 1975.
455
3. Conservation and Management of Biodiversity
Figure 29.5 Percentage of Udvardy province protected
2000 4000 6000
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Figure 29.6 Marine and coastal sites
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and Management of Biodivers:
3. Conservation
Figure 29.7 Mountainous sites
0009 O00rF O00 O
Tables 29.4 and 29.5 seem to suggest a correlation between
population pressure on an ecosystem or its economic
importance and the extent of its coverage. Thus, regions
such as mixed mountain systems or mixed island systems,
which are frequently not intensively developed, both have
extensive coverage by protected areas; temperate
grasslands, however, which are typically heavily used by
man are poorly represented. These results illustrate the fact
that socio-economic and political factors, not conservation
priorities, are often the most important considerations in the
establishing and siting of protected areas. Thus, in many
countries protected areas are established in those regions
which are the least economically valuable and with less
regard to ensuring a balanced representation of the
country’s ecosystems. This raises a number of concerns
about the ability of protected area networks alone to protect
biological diversity adequately and comprehensively.
Table 29.6 shows the present state of each country’s
protected area network and gives figures for each IUCN
Category. This table shows that, despite a global coverage
of protected areas of 5.19%, there is considerable variation
between continents. To simplify comparison, Categories I-V
have been divided into two groups: totally protected areas
with no extractive use (Categories I, II and II), and
partially protected areas with local sustainable extractive
use (Categories IV and V). Substantial variations can be
seen between countries. Fig. 29.8 illustrates the percentage
of area which is protected on a country by country basis.
The protected area network for most countries covers less
than 5% of the surface area. The map in Fig. 29.9 shows
the period during which the greatest growth occurred for
each county.
Management and funding
Any analysis of protected areas and the role they play in
conservation of biodiversity is of limited value unless there
is some assessment as to whether the protected areas are
managed properly. Developing objective indicators to
measure the degree of implementation is difficult, as proper
management of a protected area is dependent on so many
factors. This is an area in which IUCN and WCMC are
working in an effort to develop reliable indicators by which
management can be accurately assessed.
At one level, effective management requires there to be the
necessary political will. One indicator of this is the
promulgation of appropriate legislation. Another
requirement is an administrative structure with sufficient
authority and resources to manage the network adequately.
Levels of funding can therefore illustrate the commitment
or priority given to the establishment and management of
protected areas and conservation of biodiversity in general.
WCMC is beginning to compile information on funding
levels for protected areas, on a country by country basis.
The information WCMC compiled to date is given in Table
29.7. It should be noted that in many instances independent
verification of the levels of funding given in this table has
not yet been made. Consequently, the figures provided are
indicative only. Comparisons between countries at this stage
could be misleading and inaccurate. These figures do show,
however, that the amount of state funding devoted to
459
Protected Areas
protected area management in affluent countries is a
different order of magnitude from that in poor countries.
Thus, the annual budget for the USA of about US$2 billion
dwarfs the typical budget of many less developed countries,
which rarely exceeds US$500,000. Despite the limitations
of the data the table indicates that many countries do devote
considerable resources to protected areas management.
INTERNATIONAL PROTECTED AREA SYSTEMS
In the field of nature conservation there are two
international conventions and one international programme
that include provision for designation of internationally
important sites in any region of the world. These are the
World Heritage Convention, the Ramsar (Wetlands)
Convention, and the UNESCO Man and the Biosphere
(MAB) Programme. While there is a wide range of other
international conventions and programmes, these cover only
regions, or small groups of countries.
Both World Heritage sites and Ramsar sites must be
nominated by a State that is party to the relevant
convention. While there is an established review procedure
for World Heritage sites (and nomination is not guarantee
of listing), all nominated Ramsar sites are placed on the
List of Wetlands of International Importance. Biosphere
reserves are nominated by the national MAB committee of
the country concerned, and are only designated following
review and acceptance by the MAB Bureau.
Each Contracting Party to the Ramsar (Wetlands)
Convention is obliged to nominate at least one wetland of
international importance. However, a country can be party
to the World Heritage Convention without having a natural
site inscribed on the List, and may participate in the MAB
programme without designating a biosphere reserve. See
Chapter 31 for an additional view on these and other
conventions.
Wetlands of International Importance (Ramsar Sites)
The Convention on Wetlands of International Importance
especially as Waterfowl Habitat was signed in Ramsar
(Iran) in 1971, and came into force in December 1975. This
convention provides a framework for international
cooperation for the conservation of wetland habitats. It
places general obligations on contracting party states
relating to the conservation of wetlands throughout their
territories, with special obligations pertaining to those
wetlands which have been designated to the ‘List of
Wetlands of International Importance’.
Each State Party is obliged to list at least one site. Wetlands
are defined by the convention as: areas of marsh, fen,
peatland or water, whether natural or artificial, permanent
or temporary, with water that is static or flowing, fresh,
brackish or salt, including areas of marine waters, the depth
of which at low tide does not exceed six metres. Fig. 29.10
shows the parties to the Ramsar convention plus the
locations of Ramsar sites around the world.
World Heritage Sites
The Convention Concerning the Protection of the World
Cultural and Natural Heritage was adopted in Paris in 1972,
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463
3. Conservation and Management of Biodiversity
Figure 29.8 Percentage of country protected
0009 O00” 0002
=
wy
uoljdafo14 UOsUIqoy
Protected Areas
Figure 29.9 Period of greatest growth
0009 O00r O00Zd
SD81D pajoe}01d ON lel
jussaid—|96| re
sto l=GL|! |
uol}safo14 UOSUIGoY
3. Conservation and Management of Biodiversity
Table 29.7 Protected areas and levels of state financing
COUNTRY AMOUNT/ MILLION YEAR NOTES SOURCE
CURRENCY us$
ASIA
Afghanistan 0.05 PRO Proposed estimate for conservation projects by the B1
1991 Directorate of Wildlife and National Parks
Bhutan 2,128,000 Nu 0.137 1988/ Budget for Northern and Southern Wildlife circles was Al
1989 US$27,300 and $1,708,000 respectively
Brunei 59.5 million BS (1.65 = 1991 Comprises Government allocation to Forestry Department A3
1US$) (4M), Brunei Museum (7.5M), Agriculture Department
(27M), Fisheries Department (4) and District Offices (17M);
proportion subsequently allocated to protected areas is
unknown
Cyprus 637,000' Cf£ 1.28! 1991 ‘Ministry of Agriculture budget for the establishment, B3
138,000? 0.28? development and conservation of protected areas
0.035% 2Ministry of Agriculture budget for park recreational
facilities
3Running expenditure for main protected area, Lara reserve
India - Project 16.2 million Rs (29.1 Rs 1982 Allocated for 11 tiger reserves covering total area of A6
Tiger = 1 USS) 15,800km?
Indonesia 39.8 1982- Estimated government allocation for 1982-92; excludes
1992 international and private sector funding
6.0 1990- Investment from private sector for tourism in protected
1991 areas
Israel 14.5 1985 Forestry department overall yearly budget, including BS
28 1990 element for forest conservation
Jordan 32,326 JD 0.12 1990 ‘Funding for the National Environmental Strategy during the B6
(in addition to (in period up to 1990 totalled US$123,798 and JD 32,636,
USS) addition to funded by IUCN and USAID
JD
currency)
Kuwait 2 million’ KD (KD 0.294 1990’ 'The government of Kuwait allocated funds for planning, B7
= 1 US$) research and assessment of the country’s desert renewable
resources
0.7-0.9 1986? *The original proposals for annual running budgets for the
million? first national park with a fully functional administration and
necessary maintenance
%Amount allocated for all park maintenance and wardening
1 million? in Kuwait
Laos 0.01 ?1991 AQ
Lebanon 0.5! 1990 'FAO assistance to Department of Forests and Natural Bs
Resources institutional strengthening
0.2? ?FAO assistance for forestry activities including
conservation
Malaysia 4.98 1991 AS
Myanmar 1.4 71991 AS
Nepal 124.3 million NRs 2.9 1990/ Represents expenditure for total protected areas network of A7
1991 10,910km?, but 84% is allocated to Royal Nepal Army
protection units
Pakistan - 93.4 million Rs 4.3 1990/ 52.5% represents recurrent costs and 47.5% capital A2
Federal units 1991 development costs (breakdown for provinces is available)
Of Rs 2 billion allocated to forestry sub-sector in 7th Five-
Year Plan (1988-1993), Rs 332 million (16.6%) is
earmarked for wildlife conservation (Sheik and Jan, n.d.)
Philippines (816,200
US$)
Saudi Arabia c.US$ 1990- NCWCD budget total per year including all aspects for B10
9.06 1991 protected areas, administration and management. The
budget for IUCN support for two years was SAR1,444,304
(US$380,000) (IUCN 1987a)
Taiwan 36.769 1989 Total budget allocation for 4 national parks covering A8&
2,416km?
Thailand 15.097 ?1991 AS
466
Protected Areas
Table 29.7 Protected areas and levels of state financing (continued)
COUNTRY AMOUNT/ MILLION YEAR NOTES SOURCE
CURRENCY US$
ASIA (continued)
Turkey 300 million TL (6222 = 1985 Annual budget expended on park management by the B12
1 US$) General Directorate of Forestry
Viet Nam 0.222 71991 A5
Yemen 2 million’ YR (YR 1988’ ‘Forestry (former YAR) budget towards conservation B13
12.05= programmes including a national tree planting campaign
US$1) 2Budget for training of forest technicians
he 1990-
92?
Former USSR
Former USSR 54.9 million Rbl 32)7* 1990 Total investment in protected areas DS
EUROPE
Bulgaria 500 million Leva 28.016 1989 Budget for nature protection D2
Czechoslovakia- 50 million Kes 1.751 1991 Budget for protected areas (national parks 30.872 million, D3
Czech Republic CHKOs 19.102 million)
Czechoslovakia- 74 million Kes 2.592 1991 Budget for protected areas (national parks 67.342 million, D3
Slovak Republic CHKOs 6.659 million)
Denmark 6 billion DKr 980.697 1988 Overall environmental expenditure. Proposals for D4
environmental expenditure for the six year period 1989-94:
Kroner 33 billion
Estonia 634,000 Rbl 0.377* 1990 Total budget for protected areas DS
France- national 89.939 million FF 16.716 1989 Annual budget, broken down into capital (FF16.738 million) D4
parks and current or ongoing (FF73.201 million)
France- nature 14.5 million FF 2.695 1989 Annual budget, broken down into capital (FF4.5 billion) and D4
reserves current or ongoing (FF10 million)
France 6.386 billion FF 1186.880 1987 Total expenditure on protecting the natural heritage D4
(including expenditure on regional parks, parks and
gardens, other green spaces, centres for nature education,
the improvement of the surroundings of monuments, the
acquisition of green forestry spaces, forestry development
and developing fishing and hunting)
Germany- 106.415 DM 67.634 1987 Total expenditure on promotion of nature protection and
North-Rhine/ million landscape preservation, including that by non-state bodies
Westphalia (including land acquisition, preservation and development
and compensation payment). Public expenditure on nature
protection areas was DM68.386 million
Greece 200 million Drs 1.110 1991 Maximum estimate of ‘budget for protected areas’ from D6
respective bodies
Iceland 134,000 £ 0.242 1982 Running costs for the three national parks D9
Italy - state 3,022 billion Lire 2.573 1988 State environmental expenditure, divided between ongoing D4
expenditure (Lire 677 billion) and capital expenditure (Lire
2,345 billion)
Italy- regions 415 billion Lire 348 1986 Total expenditure on nature conservation in all regions and D4
and provinces provinces (from a total of Lire 3,026 billion on
environmental expenditure)
Latvia 1.188 million Rbl 0.708* 1990 Budget for protected areas DS
Lithuania 248,000 Rbl 0.148* 1990 Budget for protected areas DS
Netherlands 41 million DFI 23.110 1990 Proposed environmental spending based on the 1990 D11
Nature Policy Plan, to be spent largely on environmental
policy on specific areas, nature development, management
and maintenance
Norway 41 million NKr 6.610 1991 Total budget on protected areas, divided into compensation D12
paid for the establishment of new areas (30 million NKr)
and management or ongoing expenditure (11 million NKr)
Poland 27.472 billion ZI 2.486 1990 Expenditure for national parks D13
467
3. Conservation and Management of Biodiversity
Table 29.7 Protected areas and levels of state financing (continued)
COUNTRY AMOUNT/ MILLION YEAR
CURRENCY uss
EUROPE (continued)
Portugal 3.195 billion Esc 22.813 1991
Spain - 21.734 billion Ptas 215.852 1987
provincial
Spain - national 2.579 billion Ptas 25.611 1987
Sweden 223 million SKr 38.541 1991
UK-Northern 4 million 13 Vi2et 1991
lreland
UK - Nature 46,032,000 £ 83.165 1990/
Conservancy 1991
Council
UK - 40 million £ 72.267 1992/
Countryside 1993
Commission
UK - Gibraltar 300,000 £ 0.542 1990
UK - Isle of Man 10,000 £ 0.018 1991
NORTH AND CENTRAL AMERICA
Canada 282.99 1991
Dominica 926,300 Ec$ 0.35 1991/
1992
Dominica 1,266,730 Ec$ 0.48 1991/
1992
Guadeloupe 16 million Fr 2.97 1991
(France)
Jamaica 4.89', 1989-
2.50? 1990
Mexico 2.55 1991
St Kitts-Nevis 0.1
USA 1,962.70 1991
SOUTH AMERICA
Bolivia 2,964,915 Bol 0.784 1988
Brazil 121,139,100 Cr$ 1.211 1990
Ecuador 50.8 million Sucr 0.250 1984
Guyana 1,452,225 G$ 6.201 1990-
2000
Peru 1.162 1990
NOTES
Budget for the Portuguese Park service, of which about
64% is for capital investment and 36% for ordinary
(ongoing) expenditure. The amount spent specifically on
protected areas is probably about 68% of the total budget
Total nature conservation expenditure by the Autonomous
regions, covering: protection of flora and fauna; prevention
of forest fires; creation, conservation and management of
forests; parks and nature reserves; game hunting and
inland fishing)
National nature conservation expenditure (defined as
above)
Government money allotted to land purchase/compensation
(SKR 140 million) and to protected areas management (SKr
83 million)
Total budget of the Countryside and Wildlife Branch of the
Department of the Environment for Northern Ireland (the
bulk of this money is used for site protection)
Total expenditure
Planned expenditure for all 11 national parks in England and
Wales (including funding from Department of Environment,
local authorities and national park authorities)
Expenditure on nature and landscape conservation
(including public gardens) prior to the designation of the
only protected area
Annual government expenditure on protected areas
Canadian Parks Service and Canadian Wildlife Service
Proposed government capital expenditure on parks and
protected areas
Recurrent expenditure for Forestry Division
Budget for the ‘Parc National de la Guadeloupe’
(administrative body for Guadeloupe’s PAs)
Forest Department expenditure in the financial year,
1989/1990:
for recurrent (forest administration and soil conservation)
for capital (forestry, watershed management and
conservation)
All federal land management agency operations, including
multiple use lands
Budget for Conservation Commission
Estimates of federal government expenditures in protected
areas (includes USFWS, WWF and NAWMP)
Budget for natural resource management as a whole not
just protected area management. The CDF receives very
little external funding
Budget for federal protected areas
Budget for the national parks system
Projected budget for conservation activities including the
development of a protected area system, as part of the
National Forestry Action Plan
Budget for the entire forest and wildlife department, the
DGFF, not just protected areas. The distribution of funds
within the DGFF is not known
468
SOURCE
D14
D4
D4
D15
D16
D17
D18
D19
D20
E1
F3
F3
F2
Fil;
Anon.
1990
E1
E1
G1
G2
G3
G4
GS
Protected Areas
Table 29.7 Protected areas and levels of state financing (continued)
AMOUNT/
CURRENCY
SOUTH AMERICA (continued)
COUNTRY
Suriname 12,000 Sf
OCEANIA
Australia 15,795,193 A$
Western Samoa 104,000 Tala
AFRICA
Algeria
Angola
Benin
Burkina Faso
Cameroon
130 million F
CFA
Central African
Rep
Central African 10 million F
Rep CFA
Chad 31 million F
CFA
Congo
Cote d'Ivoire
Egypt 10 million’ E£
Equatorial
Guinea
Ethiopia
Gabon
Gambia 370,820 D
1,708,200 D
Ghana
MILLION YEAR
US$
0.007 1967
12.22 1988-
1989
0.043257 1990
0.25 1990-
1992
<0.02 1991
0.034 1991
0.5 1991
0.409 1991
0.482
0.037
0.1 1991
0.036 1991
1.321 1991
(3.3 = 1 1989-
US$) 1992
0.25 1991
0.424 1989
0.041 1990/
1991
0.19 1990/
1991
1.05 1991
NOTES
Budget for the nature protection department of the Forest
Service
Figure is for revenue, not expenditure
Proposed budget
Budget allocated to a single management plan preparation
by the BNEF as a trail project for future reorganisation of
other existing protected areas in the country
Annual personnel costs US$ < 20,000
Minimum possible figure
Annual personnel costs US$500,000
Recurrent budget (excl. personnel) US$388
Annual personnel costs (conservators and game guards
only) US$270,000
Recurrent budget (excl. personnel) US$33,000
Capital budget US$105,800
The National Centre for the Protection and Management of
Fauna (Centre National pour la Protection et
l’‘Aménagement de la Faune) is a self-financing
organisation, the budget being funded by hunting and ivory
taxes
The budget for the Ministry of Water, Forests, Hunting,
Fishing and Tourism
Budget does not include salaries
Minimum possible figure
Recurrent budget (excl. personnel) US$35,900 (1990
figure)
Annual personnel costs US$953,571
Recurrent budget (excl. personnel) US$117,857
Capital budget US$250,000
'Main government protected area initiative ‘Ras Mohammed
National Park project’ from initial phases in 1989 to project
completion. From 1989-1991 ECU 750,000 have been
budgeted, funded with technical support from the EC
The Directorate of Forestry combines four sections, one of
which is the Hunting and Protected Areas Service (Servicio
de Caza y Areas Protegidas), although this has no
personnel, vehicles or equipment
Annual personnel costs US$97,948
Recurrent budget (excl. personnel) US$153,623
Capital budget US$251,171
Under 1987-90 development plans approximately 11% of
the forestry budget (CFA 312 million) was budgeted for
forest conservation (3)
Department of Wildlife Conservation (estimated)
Forestry Department (estimated)
Annual personnel costs US$636,255
Recurrent budget (excl. personnel) US$55,327
Capital budget US$360,000 (1989)
469
SOURCE
G6
H1
B2
K1
K1
K1
K1
K4
K1, KS
K1
K1
B4
K2
K1
K1
J2
K1
3. Conservation and Management of Biodiversity
Table 29.7 Protected areas and levels of state financing (continued)
COUNTRY AMOUNT/ MILLION YEAR NOTES SOURCE
CURRENCY us$
AFRICA {continued}
Kenya 18.2 1989 Annual personnel costs US$10,000,000 (1989) K1
Recurrent budget (excl. personnel) US$84,200,000 (1989)
Capital budget US$0)
Malawi 0.456 1991
Mauritius In 1983/4 the Forest Service accounted for 0.3% of the J3
total national budget and 13.5% of the Ministry of
Agriculture, Fisheries and Natural Resources’ budget
Morocco 5 million DH 0.64 1991 In 1988, 8% of the Water and Forests budget went BS
towards parks and reserves administration and
management
In 1991, Water and Forests maintenance costs including
salaries, equipment and information dissemination
Mozambique 0.448 1986 Total annual allocation of resources available to J4
government agencies for conservation
Namibia 0.350 1991 Figure refers to Etosha NP K1
Annual personnel costs US$280,000
Recurrent budget (excl. personnel) US$70,000
Niger 1.423 1991 Annual personnel costs US$179,116 K1
Recurrent budget (excl. personnel) US$71,326
Capital budget US$14,285
Nigeria 1.66 1991 K1
Rwanda 4.73 1990 All figures for first 9 months of 1990 K1
Annual personnel costs US$1,830,000
Recurrent budget (excl. personnel) US$2,900,000
Capital budget US$4,730,000
Senegal 0.624 1991 figure refers to Niokolo Koba NP K1
Annual personnel costs US$534,857
Recurrent budget (excl. personnel) US$89,129
Sierra Leone 0.005 1991 Annual personnel costs US$4,591 K1
Recurrent budget (excl. personnel US$388)
South Africa- 0.012 1986 Total annual allocation of resources available to J4
Natal province government agencies for conservation
South Africa 3.009 1991 Figure refers to Kruger and Addo NPs; elephant budget only K1
St Helena (UK) 7,000 e 0.012 1983/ Funding (through Project-UK) from WWF/UK, ODA, FFPS L1
1984 and the British Council for conservation purposes
Sudan 1.0 1986 National Forestry Corporation budget is >68 million K1
Sudanese pounds p.a. (US$12.5 million)
(M.E.A.A.Ali, pers. comm., 1991)
Tanzania 3.478 1991 Total earnings of the Wildlife Division for 1990/1 were J5
TShs 591,676,500 (approx. US$2,572,506)
Central government returns are approx. US$1.1 million
Tanzania national Parks (TANAPA) earned >US$3.5 million
in 1990/1
Ngorongoro Conservation Area Authority earned
> US$1.84 million in 1990/1 (WD, 1991)
Togo 0.580 1990 Annual personnel costs US$405,828 (1990) K1
Recurrent budget (excl. personnel) US$10,521 (1990)
Capital budget US$20,536 (1990)
Tunisia 500,000 TD (0.9 TD 1991 ‘Maximum maintenance budget available for the key NP, B11, B14
= 1 US$) Ichkeul by the Government
2German DM 20 million made available for Ichkeul park
management via the KfW Bank, Frankfurt (1991)
Uganda 1.846 1990 K1
Zaire 1.002 1990 Annual personnel costs US$1,000,000 (1990)
Recurrent budget (excl. personnel) US$2,000 (1990)
Zimbabwe 0.009 1986 Total annual allocation of resources available to J4
government agencies for conservation
Notes: * Converted figures area based on a commercial exchange rate of US$1 = 1.6787 roubles. The free market exchange rate of US $1 = 80
roubles gives a better approximation of purchasing power.
470
Protected Areas
Figure 29.10 Wetlands of International Importance (Ramsar sites)
Uuol}dafo14 UOSUIqoY
0009 O000rv 00072
eoueywoduy
JBuoneusezu] yo spuejop,
$0 1SI] 94} UO pepnyjdu! seaqis
uonusAuod
(spuepeya) seswey 042
0} seed Buloesuo5D
471
3. Conservation and Management of Biodiversity
and came into force in December 1975. The convention
provides for the designation of areas of ‘outstanding
universal value’ as World Heritage Sites, with the principal
aim of fostering international cooperation in safeguarding
these important areas. Sites, which must be nominated by
the signatory nation responsible, are evaluated for their
world heritage quality before being inscribed by the
international World Heritage Committee. Only natural sites,
and those with mixed natural and cultural aspects are
considered in this publication.
Article 2 of the World Heritage Convention considers as
natural heritage: natural features consisting of physical and
biological formations or groups of such formations which
are of outstanding universal value from the aesthetic or
scientific point of view; geological or physiographical
formations and precisely delineated areas which constitute
the habitat of threatened species of animals and plants of
outstanding universal value from the point of view of
science or conservation; and natural sites or precisely
delineated areas of outstanding universal value from the
point of view of science, conservation or natural beauty.
Criteria for inclusion in the list are published by UNESCO.
The map in Fig. 29.11 shows the location of each World
Heritage Site plus the countries that are party to the
convention.
Biosphere Reserves
The establishment of biosphere reserves is not covered by
a specific convention, but is part of an international
scientific programme, the UNESCO Man and the Biosphere
(MAB) Programme. The objectives of the network of
biosphere reserves, and the characteristics which biosphere
reserves might display, are identified in various UNESCO-
MAB documents, including the Action Plan for Biosphere
Reserves.
Biosphere Reserves differ from the preceding types of site
in that they are not exclusively designated to protect unique
areas or important wetlands, but for a range of objectives
which include research, monitoring, training and
demonstration, as well as conservation. In most cases the
human component is vital to the functioning of the
biosphere reserve, which does not necessarily hold for
either World Heritage or Ramsar sites. See Fig. 29.12 for
the location of Biosphere Reserves. For this map only the
green tint indicates those countries which have one or more
biosphere reserves.
Table 29.8 provides summary statistics on these three
international protection systems.
References
Miller, K.R. 1980. Planning National Parks for Ecodevelopment,
Center for Strategic Wildland Management Studies, Ann Arbor.
TUCN, 1984. Categories and criteria for protected areas. In: McNeely,
J.A. and Miller, K.R. (Eds), National Parks, Conservation, and
Development. The role of protected areas in sustaining society.
Smithsonian Institution Press, Washington. Pp.47-53.
TUCN/UNEP 1986. Managing Protected Areas in the Tropics. TUCN,
Gland.
IUCN 1990. 1990 United Nations List of National Parks and Protected
Areas. TUCN, Gland. 284pp.
472
Udvardy, M.D.F. 1975. A Classification of the Biogeographical
Provinces of the World. UCN Occasional Paper No.18. Morges.
Sources for Table 29.7
Al Blower, J.H. 1989. Nature Conservation in Northen and
Central Bhutan. FAO, Rome. 48pp.
Malik, M.M. 1990. Management status of protected areas in
Pakistan. Paper presented at Regional Expert Consultation on
Management of Protected Areas in the Asia-Pacific Region.
FAO Regional Office for Asia and the Pacific, Bangkok,
10-14 December 1990. 40pp.
Othman, M. and Ramos, V.J.A. 1991. National report on
national parks and protected areas in Brunei Darussalam.
Presented to 36th Working Session of TUCN CNPPA,
Bangkok, 2-4 December. 66pp.
Sheik, M.I. and Jan, A. undated. Role of forests and forestry in
national conservation strategy of Pakistan. Draft for comment.
National Conservation Strategy Secretariat, Islamabad. 86pp.
World Bank 1991. Conserving biological diversity: a strategy
for protected areas - Asia region. Preliminary draft. 57pp.
Government of India
B. Upreti, pers. comm.
Taiwan Parks, 1984
Information provided at the 36th IUCN CNPPA Working
Session, Bangkok, 2-4 December
MacPherson, N. 1991. Opportunities improved
environmental management in Afghanistan. Report of an
TUCN mission under contract to the Office for the
Coordination of United Nations Humanitarian and Economic
Assistance Programmes relating to Afghanistan. 66pp.
Anon. 1990. Algeria, watershed management and forestry
project, conservation of nature. World development indicators
on the environment. The World Bank, Washington DC. 10pp.
Antoniou, in litt., 1991
EEAA 1991. Protected areas in the Arab Republic of Egypt.
Paper presented at the Third Man and Biosphere Meeting on
Mediterranean Biosphere Reserves and the First TUCN-
CNPPA meeting for the Middle East and North Africa, 14-19
October 1991, Tunis, Tunisia. 18pp.
Anon. 1990. National report on forestry in Israel. Report for the
Tenth World Forestry Congress by the Land Development
Authority, Kiryat-Hayim. 19pp.
McEachern, J. 1990. Report on project activities, National
Environment Strategy - Jordan September 1989 through
January 1990. National Environment Strategy, c/o Department
of Environment. IUCN Project Office, 31 January 1990.
Alsdirawi, F. 1991. Protected areas in the state of Kuwait.
Caracas Action Plan paper presented at the Third Man and
Biosphere Meeting on Biosphere Reserves in the
Mediterranean and the First IUCN-CNPPA Workshop on
Protected Areas in the North Africa-Middle East Region, 14-
19 October 1991, Tunis. 10pp.
Child, in litt., 1990
Eaux et Foréts 1991. Rapport sur les aires protégées au Maroc.
Paper presented at the Third Man and Biosphere Meeting on
Biosphere Reserves in the Mediterranean and the First IUCN-
CNPPA Workshop on Protected Areas in the North Africa-
Middle East Region, 14-19 October 1991, Tunis. 18pp.
B10 Abuzinada, A.H. and Child, G. 1991 Developing a system of
protected areas in Saudi Arabia. National Commission for
Wildlife Conservation and Development, Riyadh. Paper
presented at the Third Man and Biosphere Meeting on
Mediterranean Biosphere Reserves and the First TUCN-
CNPPA meeting for the Middle East and North Africa, 14-19
October 1991, Tunis. 16pp.
Bel Hadj Kacem, S. 1985. La conservation de la faune et de la
flore sauvages en Tunisie. Séminaire sur la conservation du
patrimoine forestier national, 30-31 octobre 1985 a l’ INPPSA
de Sidi-Thabet.
B12 General Directorate of Forestry 1987. Forestry in Turkey.
General Directorate of Forestry, Ministry of
Agriculture,Forest and Rural Affairs, Ankara.
A4
for
B2
B3
BS
B6
B7
B8
B9
Bil
Protected Areas
Figure 29.11 World Heritage Sites
0009 000% 000 O
C=
Wu
31S! eBe98} Pop, ey) UO
paquosul sezis je1n}jno
/jeanyeu pexiw pue jeinjeyy
uonueAuod
eBeW9aH POA eYyr
0} sonued Bunoejuo5
uoldefo14 UoSsUIgoY
473
3. Conservation and Management of Biodiversity
Figure 29.12 Biosphere Reserves
0009 O00rF 0002
Cz a
seAiesel o1oyudsoig
SeAsesel es9ydsoiq
yum seujunod
uol}efoi4 UOSUIqoYy
474
Table 29.8 International protection systems
Protected Areas
WORLD HERITAGE SITES BIOSPHERE RESERVES RAMSAR WETLANDS
DATE NO NO AREA (ha) DATE NO AREA (ha)
WORLD 95 (8) 320 161,944,969 538 32,336,169
ASIA 13 (2) 38 12,885,459 40 «1,354,493
Afghanistan March 1979 0 > = === = =
Bahrain May 1991 = = = == = =
Bangladesh August 1983 0 = = ==> = =
Bhutan === = = = === = =
British Indian Ocean Territory (see UK) () = = (see UK) tt) 0
Brunei ==5 = = = === = =
Cambodia November 1991 = = = === = =
China December 1985 1 (1) 8 1,966,722 --- - -
Cyprus August 1975 ) = = === - =
Hong Kong (see UK) o = = (see UK) ) )
India November 1977 5 = = October 1981 6 192,973
Indonesia July 1989 2 6 1,482,400 ==> = -
Iran, Islamic Rep February 1975 0 9 2,609,731 June 1975 18 1,087,550
lraq March 1974 0 = = === = -
Israel === = = = ———— = =
Japan === = 4 116,000 June 1980 3 9,892
Jordan May 1975 0 = = January 1977 1 7,372
Korea, Dem People’s Rep === = 1 132,000 == = =
Korea, Rep September 1988 te) 1 37,430 ==> = =
Kuwait = = = = =5= = =
Laos March 1987 0 = = === - -
Lebanon February 1983 0 = — -=-- - =
Malaysia December 1988 0 - - --- - -
Maldives May 1986 0 = = ==> = =
Mongolia _ February 1990 () 1 5,300,000 SS= = =
Myanmar ==> = > = SSS = =
Nepal June 1978 2 = - December 1987 1 17,500
Oman October 1981 to) = = =--- = =
Pakistan July 1976 () 1 31,355 July 1976 9 20,990
Philippines September 1985 0 2 1,174,345 = = =
Qatar September 1984 0 - - --- - -
Saudi Arabia August 1978 0 = ~ =-- = =
Singapore --- - - = --- - -
Sri Lanka June 1980 1 2 9,376 June 1990 1 6,216
Syria August 1975 te) = = === = =
Taiwan — - = = --- - -
Thailand September 1987 1 3 26,100 ——— - -
Turkey March 1983 1 (1) = = == - -
United Arab Emirates =S= = = = ==S = =
Viet Nam October 1987 i) = = September 1988 1 12,000
Yemen Arab Rep January 1984 # ty) - = --- = -
USSR (former) oO 20 =: 10,891,366 12: 2,987,185
Byelorussian SSR October 1988 0 1 76,201 (see USSR) = -
Ukrainian SSR October 1988 te) 2 120,655 (see USSR) = =
USSR October 1988 0 17 10,694,510 October 1976 12 2,987,185
EUROPE 11 @&) 91 4,787,243 328 Ǥ-3,782,517
Albania July 1989 0 = - --- - =
Andorra --- - - - --- - -
Austria === - 4 27,600 December 1982 5 102,369
Belgium =--- - - - March 1986 6 9,607
Bulgaria March 1974 2 17 39,922 September 1975 4 2,097
Czechoslovakia SS = 6 364,170 July 1990 8 16,958
Denmark July 1979 0 = = September 1977 27 734,468
Faeroe Islands (see Denmark) 0 - - (see Denmark) 0 0
Finland March 1987 (0) = - May 1974 11 101,343
France June 1975 1 (1) 6 575,583 October 1986 8 422,585 __
Germany August 1976 * te) 9 701,849 February 1976 ** 29 360,894
Greece July 1981 0 (2) 2 8,840 August 1975 11 107,400
Hun: July 1985 0 5 128,884 April 1979 13 110,389
Iceland --- - - = December 1977 2 57,500
Ireland September 1991 = 2 8,808 November 1984 21 12,562
Italy June 1978 0 3 3,798 December 1976 46 56,950
Liechtenstein === = = = December 1991 1 90
Luxembourg September 1983 (*) - - === = -
Malta November 1978 ) - - September 1988 1 1
Monaco November 1978 () = = = = =
Netherlands --- = 1 260,000 May 1980 1 306,348
Norway May 1977 0 1 1,555,000 July 1974 14 16,256
Polan June 1976 1 4 25,836 November 1977 5 7,090
Portugal September 1980 t) 1 395 November 1980 2 30,563
Romania May 1990 1 3 41,213 September 1991 1 647,000
San Marino a - - = ==- = =
Spain May 1982 1 10 537,717 May 1982 17 98,887
Sweden January 1985 () 1 96,500 December 1974 30 382,750
Switzerland September 1975 0 1 16,870 January 1976 8 7,049
United Kingdom May 1984 2 13 44,258 January 1976 45 173,257
Vatican City October 1982 0 = = === = =
Yugoslavia May 1975 3 (2) 2 350,000 March 1977 2 18,094
475
3. Conservation and Management of Biodiversity
Table 29.8 International protection systems (continued)
WORLD HERITAGE SITES BIOSPHERE RESERVES RAMSAR WETLANDS
DATE NO NO AREA (ha) DATE NO AREA (ha)
NORTH AND CENTRAL AMERICA 22 69 94,624,670 63 15,368,626
Anguilla (see UK) 0 - — (see UK) (1) 0
Antigua and Barbuda November 1983 0 = = === = =
Aruba (see Netherlands) 0 = - (see Netherlands) 1 70
Bahamas S== = = = ==> - -
Barbados == = = = =——= = -
Belize --- = - = =—= - -
Bermuda (see UK) 0 = = (see UK) 0
Canada July 1976 6 6 1,049,978 January 1981 30 12,937,549
Cayman Islands (see UK) tt) = - (see UK) i) i)
Costa Rica August 1977 1 2 728,955 December 1991 2 29,769
Cuba March 1981 ts) 4 323,600 a = =
Dominica === = = = SS = +
Dominican Rep February 1985 [) - = ==- = -
El Salvador October 1991 - - - --- - -
Greenland S== = 1 70,000,000 September 1977 11 1,044,500
Grenada I = = =--- - =
Guadeloupe (see France) 0 = = (see France) 0
Guatemala January 1979 1 1 1,000,000 June 1990 1 48,372
Haiti January 1980 (0) - - --- - -
Honduras June 1979 1 1 500,000 S== = =
Jamaica June 1983 0 = = === = -
Martinique Sr France) te) = = (see France) 0 ()
Mexico ebruary 1984 1 6 1,288,454 July 1986 1 47,480
Montserrat === = = = === = =
Netherlands Antilles (see Netherlands) () = = (see Netherlands) 5. Saee2010
Nicaragua December 1979 0 - - --- - -
Panama March 1978 2 1 597,000 November 1990 1 80,765
Puerto Rico SSS = 2 15,346 --- = =
St Kitts and Nevis July 1986 () = = a = -
St Lucia October 1991 = = = =-- = -
St Vincent and the Grenadines --- = = = ——— = =
Trinidad and Tobago --- = - = --- - -
Turks and Caicos Islands (see UK) 0 = = (see UK) 1 37,270
United States December 1973 10 44 19,115,210 December 1986 10 = 1,140,841
Virgin Islands (British) (see UK) te) = - (see UK) () ()
Virgin Islands (US) (see USA) t+) 1 6,127 (see USA) te) 0
SOUTH AMERICA 9 26 = 13,781,071 7 322,085
Argentina August 1978 2 5 2,409,980 === = =
Bolivia October 1976 0 3 435,000 June 1990 1 5,240
Brazil September 1977 1 2 1,862,100 =—-=- = =
Chile February 1980 te) 7 2,406,633 July 1981 1 4,877
Colombia May 1983 0 3 2,514,375 === = =
Ecuador June 1975 2 2 1,446,244 September 1990 2 90,000
French Guiana (see France) 0 - - (see France) 0 0
Guyana June 1977 0 = - --- - -
Paraguay April 1988 0 = - --- - -
Peru February 1982 4 3 2,506,739 === = =
Suriname === = = = March 1985 1 12,000
Uruguay March 1989 (0) 1 200,000 May 1984 1 200,000
Venezuela === = = = November 1988 1 9,
OCEANIA 12 13 4,745,223 45 4,515,961
American Samoa (see USA) 0 - = (see USA) 0 0
Australia August 1974 9 12 4,743,223 May 1974 40 4,477,862
Cook Islands (see New Zealand) 0 = = (see New Zealand) 0 0
Fiji --- - - - --- - -
French Polynesia == = 1 2,000 === = =
Guam (see USA) i) - - (see USA) t+) 0
Kiribati --- - - - --- - -
Marshall Islands --- - - - SSS = =
Micronesia, Federated States of === = = = aS = =
Nauru === = = = S=5 = =
New Caledonia (see France) 0 = = (see France) 0 te)
New Zealand November 1984 2 - = August 1976 5 38,099
Niue (see New Zealand) 0 = - (see New Zealand) (0) 0
North Marianas Islands --- - - - --- - =
Palau === = = = == - = =
Papua New Guinea =a = = = === = =
Pitcairn Island (see UK) 1 - = (see UK) 0 0
Solomon Islands --- - - = === = a
Tokelau (see New Zealand) 0 = = (see New Zealand) 0 0
Tonga SS5 = = = SSS = =
Tuvalu =—=— - - - --- = =
Vanuatu ss = - = =—=-- = =
Wallis and Futuna Islands (see UK) 0 = = (see UK) t) i)
Western Samoa === = = = SS = =
ANTARCTICA oO 0 o 0 0
Antarctica Sam = = = === = =
Falkland Islands (Malvinas) (see UK) (0) - - (see UK) 0 i!)
French Southern Territories (see France) te) = = (see France) 0 tt)
476
Protected Areas
Table 29.8 International protection systems (continued)
WORLD HERITAGE SITES BIOSPHERE RESERVES RAMSAR WETLANDS
DATE NO NO AREA (ha) DATE NO AREA (ha)
AFRICA 28 (1) 43 20,229,937 43 4,005,302
Algeria June 1974 1 2 7,276,438 November 1983 2 4,900
Angola =>= = = = == = =
Benin June 1982 0 1 880,000 -——= - -
Botswana — = = = === = =
Burkina Faso April 1987 i} 1 16,300 June 1990 3 296,300
Burundi May 1982 1) = = ees = =
Cameroon December 1982 1 3 850,000 === = -
Cape Verde April 1988 ts) = = === ~ =
Central African Rep December 1980 1 2 1,640,200 === = -
Chad --- - - = June 1990 1 195,000
Comoros SSS = = = a - -
Congo December 1987 0 2 246,000 ==> - -
Cote d'Ivoire January 1981 3 2 1,500,000 --- - -
Djibouti et = a = === = -
Egypt February 1974 0 1 1,000 September 1988 2 105,700
Equatorial Guinea === = = = === = -
Ethiopia July 1977 1 - = --- - -
Gabon December 1986 (0) 1 15,000 December 1986 3 1,080,000
Gambia July 1987 0 = = --- - -
Ghana July 1975 te) 1 7,770 February 1988 1 7,260
Guinea March 1979 1 2 133,300 === = =
Guinea-Bissau ==> = = = May 1990 1 39,098
Kenya June 1991 = 5 851,359 June 1990 1 18,800
Lesotho === = = = === = -
Liberia = = = = Sa = -
Libya October 1978 0 = = --- - -
Madagascar July 1983 1 1 140,000 === - -
Malawi January 1982 1 = = --- - -
Mali April 1977 Oo (1) 1 771,000 May 1987 3 162,000
Mauritania March 1981 1 = = October 1982 1 1,173,000
Mauritius sss = 1 3,594 === = -
Mayotte (see France) 0 = - (see France) 0 (0)
Morocco October 1975 (0) = = June 1980 4 10,580
Mozambique, People’s Rep November 1982 0 - - --- - -
Namibia = = = = === = =
Niger December 1974 1 - - April 1987 1 220,000
Nigeria October 1974 ts) 1 460 === - =
Reunion (see France) te) = os (see France) 0 0
Rwanda SS = 1 15,065 === = =
Saint Helena (see UK) 0 = = (see UK) 0 te)
Sao Tome and Principe === = = - ses - -
Senegal February 1976 2 3 1,093,756 July 1977 4 99,720
Seychelles April 1980 2 = = === = -
Sierra Leone ==> = - - --- -
Somalia =-- = = = --- - -
South Africa =>= = = = March 1975 12 232,344
Sudan June 1974 0 2 1,900,970 === = =
Swaziland === = = = --- - -
Tanzania August 1977 4 2 2,337,600 --- - =
Togo =-- - - - --- - -
Tunisia March 1975 1 4 32,425 November 1980 1 12,600
Uganda November 1987 0 1 220,000 March 1988 1 15,000
Western Sahara =—== - - - --- - -
Zaire September 1974 4 3 297,700 --- - =
Zambia June 1984 1 = = December 1991 2 333,000
Zimbabwe August 1982 2 = = -=- - -
Source: WCMC.
Notes: Dates are date of accession or ratification. The extra numbers in parenthesis in the World Heritage section, refer to mixed natural/cultural sites
inscribed on the list of World Heritage on the basis of beauty resulting from the man/nature interaction, rather than natural features alone. */** The former
German Democratic Republic signed the World Heritage convention in December 1988, and the Ramsar convention in July 1978. # The former People’s
Democratic Republic of Yemen signed the World Heritage convention in October 1980.
B13 FAO, in litt., 1991 D6 Kassioumis, K. 1991. Response to regional review
B14 Bel Hadj Kacem, S. 1991. Liste des parcs nationaux et aires questionnaire.
protégées - Tunisie 1991. Direction Générale des Foréts, D9 IUCN, in litt., 1982
Ministére de l’Agriculture, Tunis. Paper presented to the D11 Ministry of Agriculture, Nature Management and Fisheries
Third Man and Biosphere Meeting on Biosphere Reserves in 1990. Nature Policy Plan of the Netherlands. The Hague.
the Mediterranean, 14-19 October 1991, Tunis. 7pp. 103pp.
D2 IUCN 1991a. Environmental Status Report: 1990. Volume Two: D12 Lein, B. and Nord-Varhaug, O. 1991. Response to regional
Albania, Bulgaria, Romania, Yugoslavia. YUCN East review questionnaire.
European Programme, Cambridge, UK. D13 Oklow, C. 1991. Response to regional review questionnaire.
D3 Kucera, B. 1991. Response to regional review questionnaire. D14 Manners Moura, R. 1991. Response to regional review
D4 Cutrera, A. 1991. European Environmental Yearbook. Institute questionnaire.
for Environmental Studies. DocTer International UK/London. D15 Larsson, T. 1991. Response to regional review questionnaire.
897pp. D16 Furphy, J.S. 1991. Response to regional review questionnaire.
DS _ Nikol’skii, A., Bolshova, L.I. and Karaseva, S.E. 1991. D17 NCC 1991. Seventeenth Report 1 April 1990-31 March 1991.
Palaearctic-USSR Regional Protected Areas Review. Paper Nature Conservancy Council, Peterborough. 126pp.
proposed for IV World Parks Congress on National Parks and D18 Phillips, A. 1991. Response to regional review questionnaire.
other Protected Areas. USSR Ministry of Natural Resources D19 Cortes, J. 1991. Response to regional review questionnaire.
Management and Environmental Protection, Moscow. D20 Pinder, N.J. 1991. Response to regional review questionnaire.
477
3. Conservation and Management of Biodiversity
El
Fl
Gl
G2
G3
G4
G5
G6
Hl
Waugh and Perez Gil, 1992. Regional Review: Nearctic.
Prepared for the IV World Parks Congress, Caracas,
Venezuela, 10-21 February 1992.
Allen, B. 1990. National park planning in Jamaica: a project in
sustainable development and conservation. Paper presented to
the Association of Caribbean Studies Conference on the
Caribbean Environment, Santo Domingo, Dominican
Republic. 22pp.
Anon. 1991. Le Parc national de la Guadeloupe. Unpublished
report. Spp.
TUCN 1992. Protected Areas of the World: a review of national
systems. Volume 4. America. Draft.
Sandoval, G.J., Reyes, J.M. and Soria, J.L. 1989. Plan de
Acci6n para el Desarrollo forestal 1990-1995. Ministerio de
Asuntos Campesinos y Agropecuarios, Subsecretaria de
Recursos Naturales Renovables y Medio Ambiente, La Paz.
98pp.
Dias, I.F.O., Gongalves, A.R., Borges, M. and Meneses, E.O.
1991. Sistema de Unidades de Conservagado Federais do
Brasil. IBAMA-DIREC-DEUC. 1 Ipp.
Cabarle, B.J., Crespi, M., Calaway, H.D., Luzuriaga, C.C.,
Rose, D. and Shores, J.N. 1989. An Assessment of Biological
Diversity and Tropical Forests for Ecuador. Prepared for US-
AID/Ecuador as an Annex to the Country Development
Strategy Statement 1989-1990. 110pp.
GFC and CIDA 1989. National Forestry Action Plan 1990-2000.
Guyana Forestry Commission and Canadian International
Development Agency, Kingston, Georgetown. 77pp.
DGFF 1991. Informe sobre progreso forestal 1988-1990 del
Peri. 17th meeting of the Latin American Forestry
Commission - COFLA, Venezuela, 1 8th-22nd February 1991.
Ministerio de Agricultura, Direccién General de Forestal y
Fauna, Lima. 22pp.
Schultz, J.P. 1968. Nature preservation in Suriname: a review
of the present situation. Suriname Forest Service, Paramaribo.
21pp.
ANPWS 1989. Annual Report 1988-89. Australian National
Parks and Wildlife Service. Canberra. 132pp.
478
Il SPREP 1989. Country review: Western Samoa. Fourth South
Pacific Conference on Nature Conservation and Protected
Areas. South Pacific Commission, Noumea, New Caledonia.
12pp.
J2._—_ Edens, J.H. 1991. Tropical Forestry Action Plan - background
paper. MNRE, Banjul and FAO, Rome. 30pp.
J3 _—‘ Forestry Service 1985. Progress report 1980-84 by the Forestry
Service of the Ministry of Agriculture, Fisheries and Natural
Resources. Forestry Service, Mauritius. 14 pp.
J4 TUCN/SSC 1990. African Elephants and Rhinos Status Survey
and Conservation Action Plan. Compiled by D.H.M.
Cummings, R.F Du Toit and S.N. Stuart. 72pp.
J5 WD 1991. Elephant Conservation Plan for Tanzania. Wildlife
Division. 147pp.
K1 Data taken from the African Elephant Conservation Review
1991, produced by the African Elephant Conservation
Coordinating Group AECCG. (Data refer to government
expenditure on Wildlife and Protected Area Management,
unless otherwise stated).
K2 MALFF 1991. Elephant Conservation Plan. Equatorial Guinea.
Ministry of Agriculture, Livestock, Fisheries and Forestry.
44pp.
K3 McShane, T.O. and McShane-Caluzi, E. 1990. Conservation
before the Crisis: a strategy for conservation in Gabon.
WWF.
K4 MEFCPT 1986. Plan quinquennal secteur chasses et faune
1986-1990. Ministére des Eaux, Foréts, Chasses, Péches et du
Tourisme. Unpublished. 12pp.
K5 MTE 1991. Plan de Conservation de |’Elephant au Tchad.
Ministére du Tourisme et de l’Environnement. 49pp. Ministry
of Water, Forests, Hunting, Fishing and Tourism (Ministére
des Eaux, Foréts, Chasses, Péches et du Tourisme MEFCPT).
Based on text prepared by Sam Johnston, maps prepared by
Joel Smith, with additional material by WCMC staff.
30. MULTILATERAL TREATIES
A multilateral treaty is an international agreement concluded
between three or more states and governed by international
law.
Existing international treaties which deal entirely or in part
with biological diversity have evolved in an uncoordinated
manner. Despite this, and the consequent gaps and
duplications in overall coverage, a handful of such treaties
have come to exert a very powerful effect on the
conservation and management of elements of biodiversity.
Perhaps foremost among these, in terms of their
sophistication and global scope, are The Convention on
International Trade in Endangered Species of Wild Fauna
and Flora (CITES), The Convention on Wetlands of
International Importance (Ramsar), and The Convention
Concerning the Protection of the World Cultural and
Natural Heritage (World Heritage). The Convention on the
Law of the Sea (UNCLOS), which is yet to enter into
force, has strong potential for enhancing marine and coastal
conservation.
The names of these major treaties are indicative of their
sectoral focus, and even if the many important regional and
species-related treaties are also considered, it is clear that
the total obligations explicit in existing treaties fall short of
the demands of an adequately comprehensive system. The
proposed Convention on Biological Diversity attempts to
meet many of these demands, and is the first treaty planned
to concentrate specifically on the conservation and use of
global biodiversity (see Chapter 35).
Text
The production of a multilateral treaty usually follows
several stages. The first involves negotiation of the text of
the treaty. This can take many years and can require
numerous meetings. The negotiation of a treaty is concluded
by the adoption of the text of a treaty. This typically takes
place when all the states participating in the negotiations
teach agreement although the need for unanimity is not
required by law. Each negotiating conference adopts its own
tules concerning voting. Adoption of a treaty does not by
itself create any obligations.
Consent
A treaty does not come into force until two or more States
consent to be bound by the treaty. The expression of such
consent is usually an entirely separate process from
adoption. Consent may be expressed by “signature,
exchange of instruments constituting a treaty, notification,
acceptance, approval or accession or by many other means
if so agreed." The permitted ways of expressing consent
and becoming a party to a treaty are always outlined in the
text of the treaty itself. Signature and ratification are the
most frequent means of expressing consent. Signature refers
479
Multilateral Treaties
to the signature of the diplomats negotiating the treaty and
is often synonymous with the adoption of the treaty.
Ratification is the need for approval of the treaty by the
head of state or the legislature. In addition to signature and
ratification, a state can also become a party to a treaty by
accession. Accession is the normal way that states who did
not participate in the negotiations become parties to the
treaty. Accession is only possible if it is provided for in the
treaty; it has the same effect as signature and ratification
combined.
Entry into force
The final stage in the production of a treaty is its entry into
force. This usually occurs when all the negotiating states
have expressed their consent to be bound by the treaty. This
may be altered by agreement and it is not uncommon for
the date at which a treaty enters into force to be delayed in
order to give parties time to adapt themselves to its
requirements. Another common variation occurs when there
are a great number of states participating in the drafting of
a treaty. In this case, to wait for every State to ratify the
treaty before it enters into force would invariably cause
excess delay, and so large multilateral treaties often enter
into force when a specified number of States have ratified.
However, when this specified number is reached, the treaty
will only be in force between those States which have
ratified it; it does not enter into force for the other States
until they in turn have ratified it.
Multilateral treaty table
Table 30.3 lists all multilateral international treaties which
have been adopted for the conservation of biodiversity.
These treaties have here been classified into three broad
groups. ‘Global treaties’ are ones which have no
requirements as to membership and are open to any country
in the world. ‘Regional treaties’ are ones which limit
membership, normally to a certain geographical region,
although in some instances other criteria are used as well.
‘Species-related treaties’ are ones which limit membership
to those countries which have some relationship with the
species which are the subject of the treaty. The scope of
these treaties varies from those which, like the Antarctic
Treaty, attempt to deal comprehensively with the
governance of an area, to those, such as the Vicuia Treaty,
which confine their scope to the conservation of one single
species.
Table 30.1 represents graphically the status and membership
of global and regional treaties; Table 30.2 covers species-
related treaties. These tables are based on information
provided to WCMC by the IUCN Environmental Law
Centre (ELC) on 1 March 1992.
3. Conservation and Management of Biodiversity
Regional conventions
Alps (Salzburg) 1991
S Pacific (Noumea)
ASEAN Agt. (Kuala Lumpur)
PA Protocol (Nairobi) «
E African Reg. (Nairobi)
SPA Protocol (Kingston) *
Carribean (Cartagena de Indias)
Central African (Libreville)
Benelux (Brussels)
Red Sea (Jeddah)
SE Pacific (Lima)
W & Cent. Africa (Abidjan)
SACEP (Colombo)
CCAMLR (Canberra)
European (Bern)
Amazon Pact (Brasilia)
Persian Gulf (Kuwait)
South Pacific (Apia)
Med. Shores (Monaco)
Protocol SPA (Geneva) +
Mediterranean (Barcelona)
Game Hunting (Yamoussoukro)
Baltic Seas (Helsinki)
Amend. (Warsaw) *
Baltic Sea and Belts (Gdansk)
Amend. 2 (Tarragona) +
Amend. 1 (Tarragona) *
SE Atlantic Living Resources (Rome)
African (Algiers)
Protocol (Madrid) +
Antarctic (Washington)
Western Hemisphere (Washington)
1986
1985
1985
1985
1990
1983
1983
1982
1982
1981
1981
1981
1980
1979
1978
1978
1976
1976
1982
1976
1976
1974
1982
1973
1985
1985
1969
1968
1991
1959
1940
Global conventions
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Multilateral Treaties
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481
3. Conservation and Management of Biodiversity
Regional conventions
Alps (Salzburg) 1991
S Pacific (Noumea) 1986
ASEAN Agt. (Kuala Lumpur) 1985
PA Protocol (Nairobi) * 1985
E African Reg. (Nairobi) 1985
SPA Protocol (Kingston) * 1990
Carribean (Cartagena de Indias) 1983
Central African (Libreville) 1983
Benelux (Brussels) 1982
Red Sea (Jeddah) 1982
SE Pacific (Lima) 1981
W & Cent. Africa (Abidjan) 1981
SACEP (Colombo) 1981
CCAMLR (Canberra) 1980
European (Bern) 1979
Amazon Pact (Brasilia) 1978
Persian Gulf (Kuwait) 1978
South Pacific (Apia) 1976
Med. Shores (Monaco) 1976
Protocol SPA (Geneva) + 1982
Mediterranean (Barcelona) 1976
Game Hunting (Yamoussoukro) 1976
Baltic Seas (Helsinki) 1974
Amend. (Warsaw) * 1982
Baltic Sea and Belts (Gdansk) 1973
Amend. 2 (Tarragona) * 1985
Amend. 1 (Tarragona) + 1985
SE Atlantic Living Resources (Rome) 1969
African (Algiers) 1968
Protocol (Madrid) * 1991
Antarctic (Washington) 1959
Western Hemisphere (Washington) 1940
Global conventions
ITTA (Geneva) 1983
UNCLOS (Montego Bay) 1982
CMS (Bonn) 1979
global and regional (continued)
CITES (Washington) 1973
.
WHC (Paris) 1972
Wetlands (Ramsar) 1971
High Seas (Geneva) 1958
Liv.Res. High Seas (Geneva) 1958
Plant (Rome) 1951
Table 30.1 Multilateral treaties
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3. Conservation and Management of Biodiversity
Others
Ivory Marketing (Lilongwe) 1991
Wadden Seals (Bonn) 1990
Vicuna (Lima) 1979
Polar Bears (Oslo) 1973
CCAS (London) 1972
S Pacific Marine (Santiago) 1952
Prot. * 1956
IWC (Washington) 1946
Plants
N American Plant Protect. (Yosemite) 1976
New Varieties (Geneva) 1961
Amend. * 1967
Plant Protect (Rome) 1956
Eur. Plant Protect. orgn. (Paris) 1951
Birds
Benelux - Birds (Brussels) 1970
Protect of Birds (Paris) 1950
Fisheries
Protocol | & Il (Noumea) 1990
Drift Net (Wellington) 1989
S Pacific (Port Moresby) 1987
E Pacific Tuna (San José) 1983
N Atlantic Salmon (Reykjavik) 1982
NEAFC (London) 1980
S Pacific Fish (Honiara) 1979
Nw Alantic (Ottawa) 1978
SE Asian Fish. (Bangkok) 1967
N Atlantic Fish. (London) 1967
Skagerrak Agt. (Copenhagen) 1968
Protocol (Paris) 1984
Atlantic Tuna (Rio de Janeiro) 1966
Fisheries (London) 1964
Salmon/Baltic Sea (Stockholm) 1962
Marine Fishing (Warsaw) 10 1962
Black Sea (Varna) 1959
Danube (Bucharest) 1958
species—related
Protocol (Tokyo) + 1978
Amend. to the Annex + 1962
N Pacific Fish (Tokyo) 1952
Protocol (Oslo) » 1959
European crustaceans (Oslo) 1952
Fish, Council - Med (Rome) 1949
Inter-Am Tuna (Washington) 1949
Table 30.2 Multilateral treaties
in force
ASIA
Afghanistan
Bahrain
Bangladesh
Bhutan
Brunei
484
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Korea, Dem People’s Rep
Korea, Rep
Kuwait
Laos
Lebanon
Malaysia
Maldives
Philippines
Saudi Arabia
Singapore
Mongolia
Qatar
Myanmar
Ne
Pakistan
Oman
United Arab Emirates
Sri Lanka
Syria
Turkey
Thailand
Taiwan
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3. Conservation and Management of Biodiversity
Others
Ivory Marketing (Lilongwe) 1991
Wadden Seals (Bonn) 1990
Vicuna (Lima) 1979
Polar Bears (Oslo) 1973
CCAS (London) 1972
S Pacific Marine (Santiago) 1952
Prot. + 1956
IWC (Washington) 1946
Plants
N American Plant Protect. (Yosemite) 1976
New Varieties (Geneva) 1961
Amend. + 1967
Plant Protect (Rome) 1956
Eur. Plant Protect. orgn. (Paris) 1951
Birds
Benelux - Birds (Brussels) 1970
Protect of Birds (Paris) 1950
Fisheries
Protocol | & Il (Noumea) 1990 2) ao
Drift Net (Wellington) 1989
S Pacific (Port Moresby) 1987
E Pacific Tuna (San José) 1983
N Atlantic Salmon (Reykjavik) 1982
NEAFC (London) 1980
S Pacific Fish (Honiara) 1979
Nw Alantic (Ottawa) 1978
SE Asian Fish. (Bangkok) 1967
N Atlantic Fish. (London) 1967
Skagerrak Agt. (Copenhagen) 1968
Protocol (Paris) 1984
Atlantic Tuna (Rio de Janeiro) 1966
Fisheries (London) 1964
Salmon/Baltic Sea (Stockholm) 1962
Marine Fishing (Warsaw) 10 1962
Black Sea (Varna) 1959
Danube (Bucharest) 1958
Protocol (Tokyo) * 1978
Amend. to the Annex + 1962
N Pacific Fish (Tokyo) 1952
Protocol (Oslo) * 1959
European crustaceans (Oslo) 1952
Fish, Council - Med (Rome) 1949
Table 30.2 Multilateral treaties: species—related (continued)
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3. Conservation and Management of Biodiversity
Table 30.3 Multilateral treaties
Global conventions
1951
1958
1958
1971
1972
1973
1979
1982
1983
Plant (Rome) - International Plant Protection Convention
Liv.Res. High Seas (Geneva) - Convention on Fishing and Conservation of the Living Resources of the High Seas
High Seas (Geneva) - Convention on the High Seas
Wetlands (Ramsar) - Convention on Wetlands of International Importance Especially as Waterfowl Habitat
WHC (Paris) - Convention concerning the Protection of the World Cultural and Natural Heritage
CITES (Washington) - Convention on International Trade in Endangered Species of Wild Fauna and Flora
CMS (Bonn) - Convention on the Conservation of Migratory Species of Wild Animals
UNCLOS (Montego Bay) - United Nations Convention on the Law of the Sea
ITTA (Geneva) - International Tropical Timber Agreement
Regional conventions
1940
1959
1991
1968
1969
1985
1985
1973
1982
1974
1976
1976
1982
1976
1976
1978
1978
1979
1980
1981
1981
1981
1982
1982
1983
1983
1990
1985
1985
1985
1986
1991
Western Hemisphere (Washington) - Convention on Nature Protection and Wildlife Preservation in the Western Hemisphere
Antarctic (Washington) - The Antarctic Treaty
Protocol (Madrid)
African (Algiers) - African Convention on the Conservation of Nature and Natural Resources
SE Atlantic Living Resources (Rome) - Convention on the Conservation of the Living Resources of the Southeast Atlantic
Amend. 1 (Tarragona) - Amendment to Article Xill (1) of the Convention on the Conservation of the Living Resources of the
Southeast Atlantic
Amend. 2 (Tarragona) - Amendment for Articles VIII, XVII, XIX, and XXI of the Convention on the Conservation of the Living
Resources of the Southeast Atlantic
Baltic Sea and Belts (Gdansk) - Convention on Fishing and Conservation of the Living Resources in the Baltic Sea and the
Belts
Amend. (Warsaw) - Amendendments to the Convention on Fishing and Conservation of the Living Resources in the Baltic
Sea and Belts
Baltic Seas (Helsinki) - Convention on the Protection of the Marine Environment of the Baltic Sea Area
Game Hunting (Yamoussoukro) - Convention on the Game Hunting Formalities Applicable to Tourists Entering Countries in
the Conseil de |’Entente
Mediterranean (Barcelona) - Convention for the Protection of the Mediterranean Sea against Pollution
Protocol SPA (Geneva) - Protocol concerning Mediterranean Specially Protected Areas
Med. Shores (Monaco) Agreement concerning the Protection of Water of the Mediterranean Shores
South Pacific (Apia) - Convention on Conservation of Nature in the South Pacific
Persian Gulf (Kuwait) - Kuwait Regional Convention for Cooperation on the Protection of the Marine Environment from
Pollution
Amazon Pact (Brasilia) - Treaty for Amazonian Cooperation
European (Bern) - Convention on the Conservation of European Wildlife and Natural Habitats
CCAMLR (Canberra) - Convention on the Conservation of Antarctic Marine Living Resources
SACEP (Colombo) - Articles of Association of the South Asia Cooperative Environment Programme
W & Cent. Africa (Abidjan) - Convention for the Cooperation in the Protection and Development of the Marine and Coastal
Environment of the West and Central African Region
SE Pacific (Lima) - Convention for the Protection of the Marine Environment and Coastal Area of the South-East Pacific
Red Sea (Jeddah) - Regional Convention for the Conservation of the Red Sea and of the Gulf of Aden Environment
Benelux (Brussels) - Benelux Convention on Nature Conservation and Landscape Protection
Central African (Libreville) - Agreement for the Cooperation and Consultation Between the Central African States for the
Conservation of Wild Fauna
Carribean (Cartagena de Indias) - Convention for the Protection and Development of the Wider Caribbean Region
SPA Protocol (Kingston) - Protocol concerning Specially Protected Areas and Wildlife to the Convention for the Protection
and Development of the Marine Environment of the Wider Caribbean Region
E African Reg. (Nairobi) - Convention for the Protection, Management and Development of the Marine and Coastal
Environment of the Eastern African Region
PA Protocol (Nairobi) - Protocol concerning Protected Areas and Wild Fauna and Flora in the Eastern African Region
ASEAN Agt. (Kuala Lumpur) - ASEAN Agreement on the Conservation of Nature and Natural Resources
S Pacific (Noumea) - Convention for the Protection of the Natural Resources and Environment of the South Pacific Region
(SPREP Convention)
Alps (Salzburg) - Convention for Protection of the Alps
Species-related conventions
s
Fisheries
1949 Inter-Am Tuna (Washington) - Convention for the Establishment of an Inter-American Tropical Tuna Commission
1949 Fish, Council - Med (Rome) - Agreement for the Establishment of a General Fisheries Council for the Mediterranean
1952 European Crustaceans (Oslo) - Agreement concerning Measures for the Protection of the Stocks of Deep Sea Prawns
(Pandalus borealis), European Lobsters (Homarus vulgaris), Norway Lobsters (Nethrops norvegicus) and Crabs (Cancer
Pagurus)
1959 Protocol (Oslo) - Protocol Amending the Agreement concerning Measures for the Protection of Stocks of Deep Sea Prawns
(Pandalus borealis), European Lobsters (Homarus vulgaris), Norway Lobsters (Nethrops norvegicus) and Crabs (Cancer
Pagurus)
1952 N Pacific Fish (Tokyo) - International Convention for the High Seas Fisheries of the North Pacific Ocean
1962 Amend. to the Annex - Ammendment to the Annex to the International Convention for the High Seas Fisheries of the North
Pacific
488
Multilateral Treaties
Table 30.3 Multilateral treaties (continued)
Species-related conventions (continued)
1978 Protocol (Tokyo) - Protocol Amending the International Convention for the High Seas Fisheries of the North Pacific Ocean
1958 Danube (Bucharest) - Convention concerning Fishing in the Waters of the Danube
1959 Black Sea (Varna) - Convention concerning Fishing in the Black Sea
1962 Marine Fishing (Warsaw) Agreement concerning Cooperation in Marine Fishing
1962 Salmon/Baltic Sea (Stockholm) - Agreement on the Protection of the Salmon in the Baltic Sea
1964 Fisheries (London) - Fisheries Convention
1966 Atlantic Tuna (Rio de Janeiro) - International Convention for Conservation of Atlantic Tunas
1984 Protocol (Paris) - Protocol relating to Modification of the International Convention for the Conservation of Atlantic Tuna
1966 Skagerrak Agt. (Copenhagen) - Agreement on Reciprocal Access in the Skagarrak and the Kattegut
1967 N Atlantic Fish. (London) - Convention on Conduct of Fishing Operations in the North Atlantic
1967 SE Asian Fish. (Bangkok) - Agreement Establishing the Southeast Asian Fisheries Development Center
1978 NW Atlantic (Ottawa) - Convention on Future Multilateral Cooperation in the Northwest Atlantic Fisheries
1979 S Pacific Fish (Honiara) - South Pacific Forum Fisheries Agency Convention
1980 NEAFC (London) - Convention on multilateral cooperation in North-east Atlantic Fisheries
1982 N Atlantic Salmon (Reykjavik) - Convention for the Conservation of Salmon in the North Atlantic Ocean
1983 E Pacific Tuna (San José) - Eastern Pacific Ocean Tuna Fishing Agreement
1987 S Pacific (Port Moresby) - South Pacific Fisheries Treaty
1989 Drift Net (Wellington) - Convention for the Protection of Fishing with Long Driftnets in the South Pacific
1990 Protocol | & Il (Noumea) - Protocol | & Il to the Convention for the Prohibition of Fishing with Long Driftnets in the South
Pacific
Birds
1950 Protect of Birds (Paris) - International Convention for the Protection of Birds
1970 Benelux - Birds (Brussels) - Benelux Convention on the Hunting and Protection of Birds
Plants
1951 Eur. Plant Protect. Orgn. (Paris) - Convention for the Establishment of the European and Mediterranean Plant Protection
Organisation
1956 Plant Protect (Rome) - Plant Protection Agreement for the Asia and Pacific Region
1967 Amend. - Amendment of the Plant Protection Agreement for the Southeast Asia and Pacific Region
1961 New Varieties (Geneva) - International Convention for the Protection of New Varities of Plants (consolidated version)
1976 N American Plant Protect. (Yosemite) - North Atlantic Plant Protection Agreement
Animals
1946 IWC (Washington) - International Convention for the Regulation of Whaling
1956 Prot. - Protocol to the International Convention for the Regulation of Whaling
1952 S Pacific Marine (Santiago) - Treaty for the Permanent Commission on Exploitation and Conservation of Marine Resources
of the South Pacific
1972 CCAS (London) - Convention for the Conservation of Antarctic Seals
1973 Polar Bears (Oslo) - Agreement on Conservation of Polar Bears
1979 Vicufa (Lima) - Convention for the Conservation and Management of the Vicufha
1990 Wadden Seals (Bonn) - Agreement on the Conservation of Seals in the Wadden Sea
1991 Ivory Marketing (Lilongwe) - Agreement for the Establishment of Southern African Centre for lvory Marketing (SACIM)
489
3. Conservation and Management of Biodiversity
31.
In general, conservation action takes place ultimately within
a policy and legal framework established by national
governments (except for those resources lying outside
sovereign territory, discussed in Chapter 33). National
efforts alone, however, will not ensure that biodiversity is
adequately managed and additional international measures,
mainly supportive in nature, are often of great importance.
Much of the world’s biological diversity is found in less
developed countries which rarely are able to devote
adequate resources to conservation management. It is
therefore desirable for more affluent countries to provide
material assistance to those less affluent, and it would
clearly be equitable if those who carry the burden of
conservation were properly rewarded by those who benefit
from it, and this requires international systems through
which costs and benefits can flow freely across national
boundaries. The approaches examined in this chapter begin
to meet this requirement.
The chapter surveys some of the international measures
which support or assist national and local efforts in
biodiversity conservation through policy or legal means.
The direct support of national measures by means of
international development aid is the subject of the next
chapter.
These measures also provide mechanisms by which the
benefits of biodiversity can be registered with those that
have responsibility for its care. This is especially important
in the case of biodiversity, many benefits of which are
global rather than national in extent. Such global benefits
include, for example, the provision of migratory bird
habitats (in the case of wetlands) or carbon fixing capacity
(in the case of forests). Local people making decisions
about local resources of this kind will place little
importance on their global benefits, but if these are ignored,
there is little incentive to maintain the resources in their
current state. Systems that allow these benefits to be
registered in the state concerned are therefore required.
The most direct means of assistance is the ‘funding
mechanism’ approach, which provides a basis for funding
domestic regimes for provision of global public goods. The
best example of such a regime is the World Heritage
Convention, as discussed below.
A second route to the provision of global benefits is a
‘mutual listing agreement’. These international agreements
confer benefits through reciprocal obligations. For example,
the Ramsar agreement on wetlands provides for the listing
of at least one protected wetland site by each signatory of
the agreement and, in effect, the agreement acts as an
international notice-board whereby each signatory agrees to
confer benefits on all others (through the conservation of a
global public good), and signals this agreement by
recording the conserved site on the official list.
A third means of providing global public goods is to
‘privatise’ them by giving people the rights to compensation
for benefits produced by their local resources. This can be
done through the creation of internationally recognised
490
INTERNATIONAL POLICY AND LEGAL ASSISTANCE
property rights in the previously unowned resource. One
example of this is the privatisation of the world’s fisheries
through the development of the Economic Exclusive Zone
instrument in international law (see Chapter 33). In the case
of biodiversity, what is required is creation of intellectual
property rights in the information value of natural genetic
material, or creation of internationally transferable rights in
natural habitats (such as rights of exploration with regard to
genetic resources).
A fourth possible mechanism by which the global benefits
of a domestic resource can be registered in the state
concerned is through ‘regulated trading’. Certain tangible
goods are closely aligned with other goods whose values are
more difficult to harness. For example, many wildlife
species are traded in international markets and generate
substantial amounts of value. In itself, a piece of ivory or
a crocodile purse does not represent a return to
biodiversity, but when these goods come from natural
habitats that also contain a wide variety of unused but
potentially useful species, then the return from the utilised
wildlife may be seen as a return to the diverse habitat.
When this is the case, it is theoretically possible to
subsidise biodiversity through regulated trading in wildlife
products. This could become an additional role of the
Convention on International Trade in Endangered Species
(CITES), which is now mainly concerned with reducing the
impact of trade on exploited species.
This chapter finally looks at the UNEP Regional Seas
Programme, as an example of an international framework
intended to promote and coordinate national actions for
conservation of the coastal and marine environment in
defined supra-national regions.
FUNDING
Many countries lack the resources to address properly the
environmental problems with which they are confronted. At
the international level, the need for financial assistance to
help less developed countries tackle such problems has long
been apparent; international development aid has been an
important response to this need. Another way in which
financial assistance has been provided is through the
framework of international conventions.
Most international environmental conventions provide at
least some limited assistance to the less affluent contracting
parties by providing for the administration of the
convention, or funding national delegates to attend the
council conference or by supporting technical studies in
relation to the implementation of the objectives of the
conventions. This type of assistance, although important for
the effectiveness of a particular convention, is of limited
effect. There are, however, a number of conventions which
establish a trust fund for the explicit purpose of providing
material assistance to biodiversity conservation.
The World Heritage Fund
The best known international environment trust fund is the
World Heritage Fund (WHF) which was established
pursuant to the 1972 World Heritage Convention (WHC).
The WHF grants financial assistance to protect cultural and
natural heritage of outstanding universal value. The fund is
administered by the World Heritage Committee (the
‘Committee’), which was established within UNESCO.
The annual budget of the WHF is approximately $2.0
million (Table 31.1). This is raised by a combination of
voluntary or compulsory contributions from the contracting
parties. Although Article 15(2) lists a number of potential
sources of funding, most important is the obligation
contained in Article 16 which stipulates that contracting
parties will contribute to the fund either compulsorily or
voluntarily one per cent of their contribution to the regular
budget of UNESCO every two years. The voluntary
contributions are in effect the same as the compulsory
contributions with respect to amount and timing; the
distinction was made because it was felt that internal
ratification procedures would be simpler in some states if
contributions were technically voluntary.
The WHC does not normally allow the World Heritage
Committee to accept contributions to be used only for a
certain programme or project. An exception can be made,
however, if the Committee has decided on the
implementation of the programme or project. The
Committee is composed of 21 members elected by the
parties to the convention. Election of Committee members
must ensure an equitable representation of the different
regions and cultures of the world.
The WHF is used to provide assistance to contracting
parties to the WHC. Any State Party to the WHC may
request international assistance for property forming part of
the world cultural or natural heritage. The request should
include a description of the contemplated operation, the
necessary work, the expected cost, the degree of urgency,
and the reasons that the requesting State cannot meet the
expenses of the project with its own resources. Before
making a decision, the Committee must carry out any
studies and consultations that it deems necessary.
Under the WHC, assistance may take the form of: studies,
provision of experts and other staff to ensure that approved
work is carried out, training of staff and specialists, supply
of equipment needed by the State, loans, and non-repayable
subsidies. Assistance under the WHC may also be granted
to national or regional centres for the training of staff and
specialists. Large-scale assistance must be preceded by
scientific, economic, and technical studies. Table 31.1
outlines expenditure of the WHF with regard to each of
these types of assistance.
The assistance provided by the WHF generally covers only
a part of the work necessary. The State benefiting from the
assistance must contribute a substantial share of the
resources for its programme or project, unless its resources
do not permit it to do so.
Despite the relatively small size of the fund’s resources it
does illustrate a number of important features. The WHC,
with 117 contracting parties, is one of the most widely
accepted international environmental conventions and this is
vital for the success of an international convention. The
491
International Policy and Legal Assistance
WHF is an important reason for this popularity as it
provides an incentive to balance the obligations of the
convention. Therefore, many states can see accession to the
WHC as not only satisfying a moral and political need but
also as providing some material benefit.
The system of raising contributions, based on the UNESCO
scale, means that both overall donors and recipients from
the WHF must contribute to the fund. This requirement is
important because it gives the WHF an international basis
and means that it is seen to be more than just another form
of aid from the developed world. It means that the World
Heritage Committee retains a greater degree of control over
the use of resources because it is not simply another form
of multilateral aid, and it is less likely to be the subject of
political manoeuvring.
The WHC itself is discussed briefly below, and its
importance for protected area systems noted in Chapter 29.
The International Oil Pollution Fund
The earliest example of this class of funding mechanism is
the International Maritime Organisation (IMO) Oil Pollution
Fund, established in 1971 pursuant to the International
Convention on the Establishment of an International Fund
for Compensation for Oil Pollution Damage. This
convention provides for a free-standing fund, that awards
additional compensation to any person suffering oil
pollution damage, to the extent that the protection offered
by its companion treaty, the 1971 International Convention
on Civil Liability for Oil Pollution Damage, is inadequate.
The Fund is administered by an Assembly, a Secretariat
headed by a Director, and an Executive Committee. The
Assembly consists of all Contracting States to the
Convention. The Assembly’s responsibilities include
deciding how to distribute available compensation. The
Assembly must meet once a year, and can hold
extraordinary sessions if requested by the Executive
Committee or at least one-third of the members of the
Assembly.
The Executive Committee consists of one-third of members
of the Assembly but of not less than seven or more than 15
members. There are 47 States which are members of the
IOPC Fund with a further 14 expected to join in the near
future. In electing the members of the Executive Committee
the Assembly must secure’ an equitable geographic
distribution on the basis of an adequate representation of
Contracting States particularly exposed to the risks of oil
pollution and of Contracting Parties having large tanker
fleets. The Executive Committee must meet at least once a
year. The primary responsibility of the Executive
Committee is approving the settlement of claims against the
IOPC Fund.
The IOPC Fund is financed by initial and annual
contributions. Initial contributions are payable when a State
becomes a Member of the IOPC Fund and is calculated on
the basis of a fixed amount per tonne of oil received the
year preceding the State’s entry to the convention. Annual
contributions are paid by any person who has received in
the relevant calendar year more than 150,000 tonnes of
3. Conservation and Management of Biodiversity
Table 31.1 World Heritage Fund accounts 1988-1991
1988 1989 1990 1991
FUNDS FUNDS FUNDS FUNDS FUNDS FUNDS FUNDS FUNDS
ALLOCATED OBLIGATED- ALLOCATED OBLIGATED- ALLOCATED OBLIGATED- ALLOCATED OBLIGATED-
BY SPENT BY SPENT BY SPENT BY SPENT
COMMITTEE COMMITTEE COMMITTEE COMMITTEE
US$ US$ US$ US$ US$ US$ US$ US$
Preparatory assistance 100,000 82,800 100,000 30,000 150,000 121,476 150,000 52,500
and regional studies
Technical co-operation 700,000 435,463 700,000 515,500 700,000 364,900 600,000 372,782
Training 500,000 384,430 500,000 278,500 550,000 411,500 500,000 208,185
Emergency assistance 200,000 30,000 100,000 - 100,000 41,785 100,000 2,568
Promotional activites 150,000 94,415 150,000 74,750 200,000 179,044 250,000 165,600
Advisory services 280,000 279,700 247,200 242,200 300,000 258,176 420,000 175,471
Travel for experts of - - - - 20,000 11,554 20,000 -
LDC’s of Committee
Temporary assistance 260,000 260,000 210,700 210,700 135,000 135,000 190,000 190,000
to secretariat
TOTAL 2,190,000 1,566,808 2,007,900 1,351,650 2,155,000 1,523,435 2,230,000 1,167,106
3% contingency funds - = - - 65,000 - 70,000 -
2,220,000 2,300,000
Source: World Heritage Committee Annual Accounts 1988, 1989, 1990 and 1991.
crude oil in a Member State. Annual contributions are
levied to meet the anticipated payments by the IOPC Fund
and the administrative expenses of the Fund during the
coming year. The levy of contributions is based on reports
of oil receipts which are submitted by Governments of
Member States. The contributions are paid by the individual
contributors directly to the IOPC Fund. Governments have
no responsibility for these payments.
The IOPC Fund establishes two types of accounts or funds.
The first is the general fund from which are paid the
administrative expenses of the funds and general claims.
The other type of fund is the major claims fund which are
established to meet any potential liability from major
incidents, such as the sinking of an oil tanker.
In October 1991 the Assembly decided to levy annual
contributions which amounted to £26.7 million. Of this,
£5.0 million was for the general fund, and £6.7 and £15.0
million were for specific major claims funds. The payments
made by the IOPC Fund vary considerably from year to
year. As a result, the level of contributions to the Fund
varies, as illustrated in Table 31.2.
Two important and unusual features of the IOPC Fund are
the method of raising the funds and the system of assessing
contributions. The method of fund-raising is the first
example where governments have allowed an international
fund to raise income directly from private individuals. This
method has a number of advantages over restricting the
source to the public sector. It is a more efficient in that it
Table 31.2 Contributions to IOPC fund
YEAR GENERAL MAJOR TOTAL
FUND CLAIMS FUNDS LEVY
ie £ £
1979 750,000 {o} 750 000
1980 800,000 9,200,000 10,000,000
1981 500,000 0 500,000
1982 600,000 260,000 860,000
1983 1,000,000 23,106,000 24,106,000
1984 ie} fe) ce)
1985 1,500,000 ie} 1,500,000
1986 1,800,000 ie} 1,800,000
1987 800,000 400,000 1,200,000
1988 2,900,000 90,000 2,990,000
1989 1,600,000 3,200,000 4,800,000
1990 500,000 ie} 500,000
1991 5,000,000 21,700,000 26,700,000
eliminates layers of unnecessary administration and it means
that politically unpalatable choices are more easily made. It
allows the fund to assume a degree of impartiality which is
desirable in the situations with it is involved. The other
unusual feature of this fund is the system of assessing the
amount of contributions which are required from year to
year. Unlike the World Heritage Convention, which is tied
to the UN scale, the IOPC fund has a potentially open-
ended method of calculation based upon what is required
for the purposes of the convention, not what governments
may be willing to give.
492
The Global Environmental Facility
Despite the advantages of private sources of income for an
international convention, governments are generally
reluctant to establish this type of funding mechanism.
Rather, there is a strong preference for restricting the
income of these funding mechanisms to public sources. An
illustration of this preference of donor governments to use
this ‘voluntary’ or public method of raising finances for
international environmental funds is the Global
Environmental Facility (GEF) which is examined in greater
detail in Chapter 32.
The GEF has been proposed as the vehicle for funding
arrangements pursuant to any new international
environmental agreements. As such it would take over the
role that the trust funds described above have been
established for. The International Bank for Reconstruction
and Development (World Bank) Draft Resolution on the
GEF contains a provision that embodies this approach: "The
Bank is authorised to enter into other agreements with
countries party to international agreements for the
protection of the global environment, international
organisations and other entities in order to administer and
manage financing for the purpose of, and on terms
consistent with, this Resolution.”
The GEF will establish a new multilateral fund under which
grant or concessional loans will be given on an additional
basis to developing countries to enable them to implement
programmes that protect the global commons. The GEF is
capitalised at $1.0 billion to spend by the end of 1993. The
fund is financed by voluntary contributions mainly from the
developed countries. The World Bank manages the GEF
and organises project selection, appraisal and supervision,
with UNDP and UNEP participation.
The GEF allocates resources to projects that have any of the
following aims: protection of the ozone layer, limitation of
greenhouse gas emissions, protection of biodiversity, or
protection of international waters. To be eligible for funding
the project must also (1) be within cost-effectiveness
guidelines to be defined; and (2) provide measurable
benefits to the implementing country’s economy that are too
low to trigger investment by the implementing country, or
provide global environmental benefits that warrant
modification of project design.
Projects that are economically viable on the basis of
domestic benefits and costs to the implementing country are
not eligible for GEF financing unless a compelling case is
made that the operation would not proceed without GEF
involvement.
The level of capital for the GEF is the largest ever allocated
to this type of mechanism. The GEF in its short history
has, however, been the subject of much controversy. The
important role played by the World Bank in its
administration is seen by some as compounding the
problems which the GEF was established to solve.
The Wetlands Conservation Fund
Conventions which have been established for some time are
now establishing funding mechanisms. One example of this
493
International Policy and Legal Assistance
is the 1971 Ramsar treaty where the conference of the
contracting parties in January 1990 (pursuant to resolution
C.4.3.) established a ‘Wetlands Conservation Fund’ to
assist countries to implement the objectives of the
convention (see Chapter 29 for details of Ramsar sites).
The fund established pursuant to this convention is to be
operated in a similar way to the WHF. On request from a
competent national authority, the fund may provide any
developing country which is a Contracting Party to the
Convention with financial support for wetland conservation
activities in one of the following fields: improving
management of sites on the Ramsar List (e.g. management
plans, emergency action); designating new sites (e.g.
surveys, delineation of boundaries); promoting wise use
(e.g. preparing requests to development agencies,
institutional development, training); regional and
promotional activities (e.g. seminars, public education,
information activities).
Developing countries which are not yet Contracting Parties
may request a grant to support activities necessary for
designating a site for the List (e.g. site identification,
delineation or mapping).
Applications to the fund are reviewed by the Standing
Committee and administered by the Bureau. A meeting of
a sub-committee of the Standing Committee, held in
Australia in December 1990, developed procedures for the
operation of the fund.
By early 1991, voluntary contributions had been received
from the Netherlands government and WWF, and had been
promised by the governments of Austria, Switzerland, UK
and the USA. Other governments which have indicated
interest include: Denmark, Finland, France, Germany,
Italy, Japan, Norway and Sweden.
The Kuwait Fund
The UN Kuwait Compensation Fund established pursuant to
Security Council Resolution 687 (1991) (The Kuwait Fund)
is an international fund which, although not established
pursuant to a convention, could be an indicator of possible
future developments in this type of mechanism. The fund is
intended to meet compensation claims resulting from the
Gulf War for, among other reasons, "environmental damage
and the depletion of natural resources ...".
On 2 May 1991 the UN Secretary-General presented a
report to the Security Council setting out his
recommendations for the establishment and administration
of the Kuwait Fund. The fund is to operate in accordance
with UN Financial Rules and Regulations. It will be
administered by a Commission, which will function under
the authority of the Security Council and be a subsidiary
organ thereof. The principal arm of the Commission will be
a 15-member Governing Council, assisted by commissioners
to be nominated by the Secretary-General and appointed by
the Governing Council, and a secretariat.
This fund has a number of unique characteristics. Although
the Secretary-General’s report did not specify the size of the
Kuwait Fund, it is expected that it will raise up to $35
3. Conservation and Management of Biodiversity
billion over the next 10 years. This figure would make the
Kuwait Fund the largest trust fund ever established.
INTERNATIONAL OBLIGATIONS: PROTECTED
AREAS
The best way to ensure the fullest possible protection of
biodiversity is to pursue its preservation in situ. This means
protecting natural habitat to the extent that the integrity of
all of its ecological functions are maintained. i
The most important mechanism used in international treaties
to protect natural habitats is the inclusion of an obligation
for the parties to establish protected areas. The paragraphs
below discuss international systems from this point of view;
sites and coverage are discussed in Chapter 29.
These obligations facilitate the protection of natural habitats
in several ways. Firstly, they are public declarations by
governments committing themselves, morally if not legally,
to protecting natural habitats. This public commitment may
then be exploited by interested parties within a State to
promote the establishment of protected areas needed to
satisfy the obligations of the convention. This can be an
effective means of overcoming government inertia,
reluctance or opposition. A prime example of this is the use
of the obligations in the World Heritage Convention by
environmental pressure groups in Australia; several new
national parks have been created despite strong opposition
within government.
International obligations are also useful because of the clear
capacity for mutual gain to be achieved by mutual
obligations regarding the protection of natural habitats.
Each State that undertakes to protect some parts of its
diverse natural resources benefits from undertakings made
by other parties. However, it is also limited for the same
reason. This is because the world’s diverse resources are
not uniformly distributed across all nations; some -have
much more and others much less of the global total.
Reciprocity in the declaration of equal amounts of protected
areas is not a sufficient basis for ensuring full protection of
the diversity that exists in those States with the greatest
shares.
The development of the ‘mutual listing’ mechanism has
evolved with the changing attitude of man towards nature.
Initially, this mechanism was incorporated into conventions
whose primary purpose was the protection of ‘important’
wildlife, by the establishment of game reserves. An early
example of this is the 1909 Convention for the Preservation
of Wild Animals, Birds and Fisheries in Africa which
‘encouraged nature reserves’. A few decades later the
intrinsic value of natural habitat itself, as something more
than the producer of game, came to be recognised. One of
the first treaties to incorporate this shift in emphasis to the
protection of natural habitat for its own sake was the 1940
Washington Convention on Nature Protection and Wildlife
Preservation in the Western Hemisphere (Western
Hemisphere Convention).
This convention became a model for many subsequent
treaties. Its operative language called upon the contracting
parties to establish various types of protected areas, and
494
then to list these with the Organisation of American States.
The four types of protected areas defined in the convention
are: National Parks, National Reserves, Nature Monuments,
and Strict Wilderness Reserves. The careful definition of
what constitutes a protected area and the provision of an
international ‘notice-board’ for making these designations
public are the essential ingredients of a listing regime.
Although the Western Hemisphere Convention was the first
to extend protection to habitat for reasons other than game
and wildlife conservation, the intended scope of the treaty
remained somewhat narrow. It provided only for the
protection of areas labelled of special significance because
of a special animal or monument.
During the 1960s the concept of what was of special
significance and therefore worthy of protection expanded to
include areas of particular biological richness and diversity,
even though the areas might not necessarily include any one
species of special significance. This development is well
illustrated by the adoption of the 1971 Convention on
Wetlands of International Importance especially as
Waterfowl Habitat (Ramsar). Wetlands had long been under
particular threat and were generally regarded as wastelands.
However, the wide range of ecosystem services rendered by
these wetlands, in the maintenance of fisheries, wildlife and
general services, came to be recognised and the result was
a protected areas convention providing for the mutual
obligation of all parties to designate protected wetlands.
As the perceived threats to protected areas have changed so
has the nature of the obligation built in to establish such
areas. Initially, the integrity of a protected area was
believed to be safeguarded by simply ensuring that activities
within the area were controlled. In the early treaties, such
as the Western Hemisphere Convention, no attention is
given to activities outside the protected area which may
have a harmful effect on its integrity; this was remedied in
later treaties. An example of this is the 1968 African
Convention on the Conservation of Nature and Natural
Resources (African Convention), which requires parties to
the convention to establish buffer zones in order to control
activities "which may have harmful consequences on the
ecosystem" within the established protected areas. By 1985
when the ASEAN Agreement on the Conservation of
Nature and Natural Resources (ASEAN Convention) was
adopted, the establishment of buffer zones had become
standard practice.
Probably the best-known example of this approach is the
UNESCO Man and the Biosphere Programme (MAB). This
programme was established to promote sustainable
utilisation of natural resources, and to protect natural
habitats from incompatible developments in the immediate
vicinity. Initiated officially in 1971, MAB was a direct
consequence of the Biosphere Conference of 1968 and the
earlier international biological programme of the
International Council of Scientific Unions. MAB became
operational in 1976, and provides for the establishment of
‘Biosphere Reserves’ of various types throughout the world.
UNESCO biosphere reserves are a special kind of protected
area that rely upon zoning (i.e. designated land-uses) to
safeguard biological diversity. In theory, a biosphere
reserve encompasses a core zone that represents one of the
earth’s major ecosystems and is large enough to permit in
situ conservation of its genetic material. These core zones
are meant to be undisturbed by human activity, except for
scientific research. Multi-use buffer zones are intended to
surround the core, and these should be managed for the
economic benefit of local populations.
Recently, the protected area approach has been extended to
protect natural habitats in the international commons,
including the High Seas, Antarctica and Outer Space. In
these instances, parties have agreed to protect natural
habitats not by establishing protected areas but by mutually
agreeing to regulate or ban certain activities in the area
concerned. This type of protection is illustrated by the
Antarctica Treaty System where under the most recent
protocol to the Antarctic Treaty the entire area is to be
declared a protected area.
The extent of the obligations created in these international
instruments can vary from the mandatory to the purely
hortatory. Most examples are intermediate. For example, in
the Ramsar Convention the obligation to protect natural
habitat is relatively generalised; Article 4(1) of that treaty
merely requires “each contracting party to promote the
conservation of wetland and waterfowl by establishing
nature reserves on wetlands". However, in order to become
a party to the convention the State must nominate at least
one area to be included in the list of significant wetland
sites. The World Heritage Convention includes more
detailed and specific obligations; Article 4 requires each
contracting party to recognise the duty of identification,
protection, and conservation of natural heritage as defined
in the convention. It goes on to require each party to "do all
it can to this end, to the utmost of its own resources and,
where appropriate, with any international assistance and
cooperation".
The benefits of careful construction of the language of
obligation are seen when attempts are made to enforce these
undertakings. The nature of the obligation created by the
World Heritage Convention has been the subject of judicial
consideration in a series of cases in Australia, where the
High Court held that the language of Articles 4 and 5S
created a binding obligation on the contracting parties to do
all they can to protect sites on the World Heritage List.
The extent to which these international obligations have led
to increased protection of natural habitats by means of
protected area establishment is difficult to assess accurately;
certainly, many such areas are now listed as World
Heritage or Ramsar sites, or as Biosphere Reserves (see
Chapter 29). It is clear that even though the effect of these
obligations may be hard to quantify, they have been an
important method of protecting the world’s biological
diversity.
INTELLECTUAL PROPERTY RIGHTS FOR
BIOTECHNOLOGY
Intellectual Property is the term used to describe the branch
of law which protects the application of thoughts, ideas and
information which are of commercial value. It thus covers
the law relating to patents, copyrights, trademarks, trade
secrets and other similar rights (Cornish, 1989).
495
International Policy and Legal Assistance
The development of the genetic resources of biodiversity is
known as biotechnology. Broadly defined, biotechnology
includes any technique that uses living organisms or parts
of organisms to make or modify products, to improve plants
or animals, or to develop microorganisms for specific uses
(Congress of the United States, Office of Technology
Assessment, 1990). Mankind has used forms of
biotechnology since the dawn of civilisation. However, it
has been the recent development of new biological
techniques (e.g., recombinant DNA, cell fusion, and
monoclonal antibody technology) which has raised
fundamental social and moral questions and created
problems in intellectual property rights.
Intellectual property protection for biotechnology is
currently in a state of flux. Whilst it used to be the case that
living organisms were largely excluded from protection,
attitudes are now changing and increasingly biotechnology
is receiving some form of protection. These changes have
largely taken place in the USA and other industrialised
countries, but as other countries wish to compete in the new
biotechnological markets, they are likely to change their
national laws in order to protect and encourage investment
in biotechnology.
There is at the moment no clear international consensus on
how biotechnology should be treated. Although bodies such
as the World Intellectual Property Organization (WIPO, the
United Nations permanent body primarily responsible for
international cooperation in intellectual property), and the
Organization for Economic Cooperation and Development
(OECD) have conducted separate studies and produced
various reports, these have only sought to make
governments more aware of the potential problems and to
offer some suggested solutions. In view of the highly
controversial nature of providing intellectual property
protection for biotechnology, it is likely that in the short
term developments will be at a national and regional level.
Intellectual property protection currently available
There are currently two main systems of protection for
biotechnology: rights in plant varieties, and patents. Both
systems provide exclusive, time-limited rights of
exploitation and are described in more detail below.
Keeping biotechnology ‘secret’ can also be a valuable form
of protection. National treatment of trade secrets is diverse,
and all attempts to harmonise trade secret laws in Europe,
for example, have failed. Most jurisdictions do provide
some form of protection against those who steal or use
others’ trade secrets unfairly. However, the problem with
this form of protection is that the secret generally becomes
public once the biotechnology is used commercially and
thus the protection is lost.
It is conceivable that the law of copyright could afford some
protection for biotechnology. Lines of genetic code are
analogous to some extent with computer program code,
which has now been incorporated into the copyright systems
of most industrialised countries. However, this route to
protection is fraught with practical and conceptual
difficulties and is generally thought to be unsuitable. There
is as yet no recorded case of biotechnologists claiming
copyright in their inventions.
3. Conservation and Management of Biodiversity
Trademarks are also unlikely to be of much use in
protecting biotechnology, though they may of course prove
important later in regard to marketing products, processes
or services. An attempt to register the name of a plant or an
animal as a trade mark is unlikely to be successful as public
policy would prevent it (in England, registrations for names
of varieties of roses have been removed from the Trade
Mark Register for lack of distinctiveness and because of the
likelihood of confusion).
Rights in plant varieties
Prior to the mid-1960s only a few countries (e.g.,
Germany, USA) gave any intellectual property protection to
plant varieties. Because of pressure from their plant
breeding industries, 10 western European countries entered
into a diplomatic process in the early-1960s which
eventually culminated in the formation of an International
Union for the Protection of New Varieties of plants
(UPOV) and the signing of a Convention (the UPOV
Convention 1961). Since that time a number of other
countries have become parties to the UPOV Convention (the
full list of 19 parties appears in Table 31.3). Amendments
were made to the UPOV Convention in 1978, principally to
facilitate the entry of the USA.
The UPOV Convention requires that each member country
must adopt national legislation to give at least 24 genera or
species protection, in accordance with the provisions of the
convention, within eight years of signing. A plant variety is
protectable ("a protectable variety") under the UPOV
system if it is distinct, uniform, stable (DUS) and satisfies
a novelty requirement. Novelty and distinctiveness equate
broadly to novelty under patent law, but are more leniently
applied in comparison to the patent rule. Satisfaction of the
DUS criteria is conducted by the national authority
responsible, usually by growing the variety over at least
two seasons. There is also an important requirement that the
variety be maintained throughout the duration of protection.
A country may apply the system to all genera or species,
but there is no obligation to do so and thus the system has
been extended only gradually. In addition, the UPOV
Convention allows national legislation to discriminate
against foreigners (including nationals of a UPOV
Convention country) under the principle of reciprocity.
Thus amongst the UPOV members there is still some
disparity in protection.
Duration of protection depends on national legislation and
on the plant species to which the variety belongs, but is
generally for 20-30 years. Grant of plant variety rights
confers certain exclusive rights on the holder, including the
exclusive right to sell the reproductive material (e.g. seed,
cuttings, whole plants) of the protected variety. However
the rights do not extend to consumption material (e.g. fruit,
wheat seed grown for milling flour). Essentially the
exclusive rights define what others may or may not do in
relation to the protected varieties.
Plant breeders were for some time dissatisfied with the
protection provided by the UPOV system. This eventually
resulted in a major diplomatic conference in March 1991,
at which the UPOV Convention was substantially revised.
The new 1991 text will provide far greater protection than
496
is afforded at present, most notably by requiring that all
member countries apply the convention to all genera and
species, by extending the exclusive rights to include
harvested material (e.g., fruit, wheat grown for milling into
flour) and, most controversially, by allowing enforcement
against farm-saved seed (where a farmer produces further
seed of the protected variety from the previous year’s crop).
However, until the national governments ratify the new
convention the system will continue to be based on the 1978
text. There will be considerable national opposition to the
strengthening of plant variety rights and thus these changes
may take years before they are implemented and may even
be superseded by greater availability of patent protection in
the meantime.
Patents for biotechnology
A patent is a grant of exclusive rights for a limited time in
Tespect of a new and, useful invention. The exact
requirements for grant of a patent, the scope of protection
it provides and its duration differs depending on national
legislation. However, generally the invention must be of
patentable subject matter, novel (new), non-obvious
(inventive), of industrial application and sufficiently
disclosed. A patent will provide a wide range of legal
rights, including the right to possess, use, transfer by sale
or gift, and to exclude others from similar rights. Duration
will be for around 20 years (although for only 17 years in
the USA). These rights are generally restricted to the
territorial jurisdiction of the country granting the patent and
thus an inventor wishing to protect his/her invention in a
number of countries will need to seek separate patents in
each of those countries. Whilst the majority of countries
provide some form of patent protection, only a few provide
patent protection for biotechnology (these include:
Australia, Bulgaria, Canada, Czechoslovakia, Hungary,
Romania, Japan, the Soviet Union and the parties to the
European Patent Convention). The reasons for this may
differ, but generally it has been because biotechnology has
been thought inappropriate for patent protection, either
because the system was originally designed for mechanical
inventions, or for technical or practical reasons, or for one
or more ethical, religious or social concerns. In all the
National Patent Offices where patents are granted for
biotechnology there is a considerable backlog of pending
applications. Even in those countries where patent
protection is provided, the type and extent of that protection
is different in nearly every national system.
It has largely been the USA which has broken new ground
in providing the possibility of patent protection for
“anything under the sun that is made by man”. Patents have
been granted for plants since 1930 in the USA, under The
Plant Patent Act. However, prior to 1980, the US Patent
Office would not grant utility patents (separate from The
Plant Patent Act) for living matter because it deemed
products of nature not to be within the terms of the utility
patent statute. That was until the landmark decision of the
US Supreme Court in Diamond v Chakrabarty (from which
the above quote is taken), which held that a particular
genetically engineered bacterium was statutory subject
matter for a utility patent. This decision has been the basis
upon which patents have been granted for higher life forms.
Subsequently it has been held that a utility patent may be
International Policy and Legal Assistance
Table 31.3. International intellectual property treaties (party states as at 1 January
1991)
PARIS UPOV MICRO PCT EPC PARIS UPOV MICRO PCT EPC
ASIA NORTH AND CENTRAL AMERICA
Bangladesh e Bahamas e
China ® Barbados ® e
Cyprus ® Canada ® e
Indonesia ° Cuba e
Iran, Islamic Rep e Dominican Republic e
Iraq e Haiti e
Israel ® e Mexico e
Japan e e e ® Trinidad and Tobago e
Jordan ® United States ®
Korea, Dem People’s Rep ® e
Korea, Rep e e e SOUTH AMERICA
Lebanon = Argentina e
Malaysia e Brazil e e
Mongolia = Suriname e
Philippines e e Uruguay e
Sri Lanka e e
Syria e AFRICA
Turkey © Algeria e
Viet Nam ® Benin e e
USSR Burkina Faso e e
Burundi e
Soviet Union’ e e e Cameroon e e
Central African Rep e e
EUROPE Chad e e
Austria e e e e Congo ® e®
Belgium e e e e e Cote d'Ivoire e
Bulgaria e e e Egypt e
Czechoslovakia e e Gabon e e
Denmark* e ° e e e Ghana ®
Finland e e e Guinea e
France e e e e e Guinea-Bissau °
Germany e e e e e Kenya e
Greece e e e Lesotho e
Hungary e e e e Libya e
Iceland e Madagascar e e
Ireland* e e Malawi ® e
Italy* e e e e e Mali e
Liechtenstein ° e e e Mauritania e °
Luxembourg e e e Mauritius e
Malta e Morocco e
Monaco e e Niger e
Netherlands* ® e e e e Nigeria e
Norway e ® ° Rwanda e
Poland e e ° Senegal ° e
Portugal * e South Africa ° e
Romania e ° Sudan e e
Spain* ° ° ° ° e Tanzania e e e e
Sweden e ° e® e e Togo e e
Switzerland e e e e e Tunisia e
United Kingdom* e e e e e Uganda e
Vatican City ® Zaire °
Yugoslavia e Zambia e
Zimbabwe e
OCEANIA
Australia e e e e
New Zealand e e
Source: World Intellectual Property Organisation 1991.
Note: * Member States of the European Community ‘ Refers to former USSR.
Key to column headings: PARIS - means the State is a member of the International Union for the Protection of Industrial Property (Paris Union),
founded by the Paris Convention for the Protection of Industrial Property, and has ratified or acceded to at least the administrative and final
provisions Articles 13 to 30) of the Stockholm Act (1967) of that Convention. UPOV - means the State is a party to the International Convention
for the Protection of New Varieties of Plants (either the 1961 version or the revised 1978 version). MICRO - means the State is a party to the
Budapest Treaty on the International Recognition of the Deposit of Microorganisms for the Purposes of Patent Procedure. PCT - means the State
is a party to the Patent Cooperation Treaty. EPC - means the State is a party to the European Patent Convention.
497
3. Conservation and Management of Biodiversity
granted for plants and a patent has been granted for an
animal. Polyploid oysters, not naturally occurring, were
held to be patentable subject matter and US Patent
No.3,736,866, was issued in respect of a "transgenic non-
human mammal all of whose germ cells and somatic cells
contain a recombinant activated oncogene sequence
introduced into the said mammal, or an ancestor of said
animal, at an embryonic stage" - popularly known as the
‘onco-mouse’.
Elsewhere, the treatment of applications for patents for
living matter is far from certain. Whilst patents are granted
in many countries for plants and microorganisms, it has
been the issue of patents for animals which has been most
controversial. Whilst it is not possible to summarise
succinctly the position in the rest of the world, it is possible
to describe the present approach of those countries which
are party to the European Patent Convention (the EPC, see
Table 31.3). The EPC is a regional arrangement entered
into by 14 European countries for the purpose of making
multiple applications for any of the member countries a
great deal easier and to introduce a common system for
patent protection. An application under the EPC is for a
European patent, or Europatent, for short. If a Europatent
is granted by the European Patent Office (EPO) it has the
same effect, and is subject to the same conditions, as a
national patent in each of the member countries designated
in the application. In other words, through a single
application a bundle of national patents can be obtained.
The EPC provides that "plant or animal varieties or
essentially biological processes for the production of plants
or animals" are excluded from patent protection (although
the exclusion is expressly stated not to apply to
microbiological processes and products). These exclusions
would appear to place unequivocal prohibition on
Europatents for macrobiotechnology. However, the EPO
has been taking an increasingly narrow view of these
exclusions, and has held that they do not exclude all plants
and animals per se, but only claims for varieties of plants
or animals and that a process is not "essentially biological"
if there has been substantial interference by man.
It is also important to note that there is currently before the
European Parliament of the European Community (EC) a
proposal for a Council Directive for harmonisation of the
legal protection provided for biotechnology in the EC. This
does not propose to amend the EPC, but the present draft
proposal would make even more opportunities available for
patenting biotechnology and thus make the EC more
attractive in terms of investment in biotechnology research.
International treaties
There are three international intellectual property treaties
which are of particular importance for the protection of
biotechnology: the Paris Convention for the Protection of
Industrial Property (the Paris Convention); the Budapest
Treaty on the International Recognition of the Deposit of
Microorganisms for the Purposes of Patent Procedure (the
Deposit Treaty) and the Patent Cooperation Treaty (PCT)
(see Table 31.3).
The Paris Convention was originally signed in 1883 by just
11 countries, but now the majority of countries who have
498
any form of intellectual property law are parties to it. The
keystone to the convention is the principle of national
treatment: an applicant from one convention country shall
have the same rights in a second convention country as a
national of that second country. The convention covers
patents and defines them so broadly that it permits
application to any of the forms of industrial patents granted
under the laws of the convention countries. The most
important practical result of the convention is that it is
possible to claim priority from an application made in a
convention country for all subsequent convention countries
within 12 months of the original filing.
The Deposit Treaty, as the full title suggests, is concerned
with the deposit of examples of microorganisms for the
purposes of patent applications. Applications for patents for
biotechnology often face considerable difficulties in
describing the nature of the invention sufficiently. The
Deposit Treaty is a vehicle for solving these problems,
primarily through the setting up of a series of International
Depository Authorities (IDA) and through the recognition
by all member countries of a deposit in a single IDA.
The PCT simplifies the process of filing patent applications
simultaneously in a number of countries. Under the PCT a
single application may be filed in one of the official
receiving offices, designating any number of PCT member
countries, which can eventually result in a national patent
being granted in each of the designated states (and/or a
Europatent). A prior-art search is performed by the
receiving office and a report sent to the applicant. The
application and report are published and the application will
then move on either to an international preliminary
examination followed by national examination, or
alternatively straight to the national examination stage.
Unfortunately, the eventual outcome is not a ‘world patent’
and there is no harmonisation patent law under the PCT
apart from the procedural aspects.
Case study: the Iguana Management Programme
The Green Iguana /guana iguana of Latin America is a
highly prized source of meat and eggs. Green Iguanas are
arboreal herbivores which can grow up to 2m in length and
can weigh as much as 6kg (about 82% of the lizard is
edible). They need about half as much food as a chicken or
rabbit to produce the same amount of meat. The species is
now widely threatened because of excess hunting and
habitat destruction.
Research into the reproductive behaviour of the Green
Iguana was begun in 1983 and resulted in development of
new management techniques for ranching. A ‘genetic brood
stock’ of adult iguanas which are larger, faster growing and
more productive has been developed. The research has
largely been the work of the Pro Iguana Verde Fundacién
(formed by Dagmar Werner in 1985). The Fundacién’s
programme for training and advice on Iguana ranching is
called the Iguana Management Programme (IMP). The IMP
is based in Costa Rica but it is intended to implement it
throughout Latin America and possibly elsewhere.
The primary purpose of the IMP is to conserve living
natural resources; its basic premise is that if farmers can
Taise iguanas as a food crop, the status of the wild species
will be improved and forest clearance might be reduced.
Farmers adopting iguana ranching would have to protect or
re-establish areas of forest to provide food for stock.
Research indicates that meat production per hectare by
iguanas is approximately three times higher than by cattle.
Income can be derived from selling iguanas and their
products (meat, eggs, leather) and products from the forest.
The new technology and expertise which have been
incorporated into an iguana ranching model are being
applied for an industrial purpose (i.e. agriculture) and are
of commercial value; they thus fall within the area of
intellectual property law as applied to biotechnology. The
biotechnological components of the ranching model are the
genetic brood stock (the Fundacion has ‘bioengineered’ an
improved stock of Green Iguanas) and the husbandry
procedures (egg laying and incubation, nutrition, disease
control, release and harvesting). These are forms of
‘original or traditional biotechnology’, as opposed to ‘new
biotechnology’ which is largely laboratory-based and
dependent upon human manipulation of genetic material.
Intellectual property rights provide the means for
compensating the Fundacidn for its efforts. The
technologies involved in the IMP are vulnerable to piracy.
Much of the work of the Fundacién is contained in the
genetic make-up of the Genetic Brood Stock. Once these
Iguanas are transferred or sold the Fundacidn loses its
direct control over the animals. In addition, the success of
the Iguana ranching model is dependent on the expertise to
use the technologies efficiently; this is information which
took years to develop but which can be pirated very easily
once a licence is purchased. The Fundacién needs to be able
to disseminate its innovations and expertise in the security
of knowing that it cannot be re-sold by pirates and that
there will be no reduction of the licensing potential. Only
internationally recognised intellectual property law can
provide these types of protection.
Because of the uncertainties of the world’s intellectual laws
with regard to biotechnology the availability of protection
for the most important components of the IMP is
questionable. At present there is widespread discrimination
against the application of intellectual property rights to
natural genetic materials and in favour of human-modified
genetic materials. This provides no incentives for
exploitation of useful genetic materials in the natural
environment, even though in developing countries natural
resources are obvious subjects for investment. However,
one important way to limit conversion of natural resources
is to ensure that fair value is paid for current uses of the
existing resource base. Intellectual property rights could be
a means of influencing developing countries to maintain and
develop diverse resources in return for the value that these
resources render to the world community.
REGULATED TRADING IN WILDLIFE PRODUCTS
Regulated trading in wildlife products has the capacity of
returning benefits to the users of natural habitats. It could
do this if the trade were regulated in such a way as to
support prices, much as is done at present with respect to
agricultural commodities, where price supports provide
499
International Policy and Legal Assistance
incentives for maintaining land in its current state, as
opposed to converting it to other purposes.
At present, there is no regulated trading mechanism of
exactly this nature. There are, however, a number of
existing international agreements which do seek to regulate
trade in wildlife products. Early examples are the Western
Hemisphere Convention and the 1950 Paris International
Convention for Protection of Birds. These simply outlined
in broad terms an obligation to control trade in wildlife
products but created little structure within which these
controls could be implemented. Both conventions
consequently became ‘sleeping treaties’. Undoubtedly the
most important and effective convention which places some
control on the economic exploitation of wildlife products
and thereby protects biological diversity is the Convention
on International Trade in Endangered Species of Wild
Fauna and Flora (CITES).
The evolution of CITES
CITES is the most widely accepted of international treaties
on the conservation of natural resources. The number of
Parties has been steadily increasing from the initial signing
of the convention in 1973 to a total of 113 in 1992 (Fig.
31.1)
The convention attempts to prevent commercial trade in
species of wildlife which are in danger of extinction and to
control the trade in species which might become so if their
trade was allowed to continue unchecked. It does this by
means of two lists of species: Appendix I contains those
species banned from international commercial trade and
Appendix II, those for which trade may take place provided
that export permits have been issued. Importing countries
are obliged to ensure that all imports of Appendix I
specimens are accompanied by correct export permits.
One of the main obligations of Parties is to submit to the
Convention Secretariat annual reports of all of their trade in
species included in the Appendices. The number of annual
reports submitted is also shown in Fig. 31.1. These data are
then compiled on a computer database and in this way it is
possible to determine the global levels of trade in each
species. At a fine level of resolution, the trade emanating
from each range state can then be compared with what is
known about the wild population in that country to enable
an estimation of whether it is sustainable or whether it
might be detrimental to its survival. At a coarser scale, the
data can show long-term trends in trade levels or trade
routes, which can be used to help in understanding and
therefore controlling the trade.
The convention covers not only live animals and plants but
also products and derivatives of the species listed. These
range from whole skins and manufactured leather products,
through ivory carvings, tortoiseshell jewellery, meat, seeds,
and feathers to medicinal products extracted from plants
such as ginseng. This causes problems for the
implementation of the Convention because it is necessary
for enforcement officers to determine not only what species
the product is derived from but also whether the species is
included in the Appendices. In order to minimise the
problems of identification, where numerous species are very
3. Conservation and Management of Biodiversity
Figure 31.1 CITES: number of Parties and annual reporting
120 a Number of Parties
Number of Parties/reports
Number submitting annual reports
1980 1981 1982 1983
similar in appearance, the whole group of organisms may
be included in Appendix II even if only a small proportion
of the individual species are in trade. This provision
accounts for the majority of species covered by CITES and
includes examples such as: all parrots, which are mainly
traded as pets; all cacti, and all orchids, which are popular
in horticulture; all cats (Felidae), used mainly for the skin
trade; and all primates, which are used for biomedical
research and as pets.
Of the large number of international environmental
conventions, CITES has probably the most detailed control
structure. It was the first international wildlife treaty to
provide for explicit obligations and international monitoring.
As originally drafted, CITES provided little in the way of
a trade regulation mechanism and was seen primarily as a
protectionist measure which would essentially stop trade in
endangered species. The convention is based on the
premise, that where endangered status can be attributed to
overuse use should be withdrawn. However, conversion of
habitat rather than over-exploitation is often the primary
threat. It could be argued in these cases that maximising
value to local resource users, through regulated trading, is
more beneficial to conservation than the elimination of that
value by prohibiting trade.
Recently, the Conference of the Parties to CITES has
moved toward recognition of this problem, by adopting a
more flexible approach, with the attempted development of
various sorts of constructive utilisation systems.
As early as 1979, the delegates from developing countries
brought the anomaly of "indirect extinction in lieu of direct
over-exploitation" to the attention of the Conference of the
Parties. In San José, Costa Rica, they argued that there
1984
500
1986
1986 1987 1988 1989 1990
Year
must be an economic benefit from the protected species to
justify protecting their habitats from development. These
concerns led to a first step towards the reform of CITES,
with the adoption of Conference Resolution 3.15 at the New
Delhi Conference of the Parties in 1981. This resolution
provides for the transfer of certain Appendix I populations
to Appendix II for the purposes of sustainable resource
management. The criteria which specify how Appendix I
species may be used in order to procure compensation for
their habitat are known as the "ranching criteria", and each
subsequent Conference of the Parties has seen a number of
such proposals for review and possible acceptance. The first
ranching proposal accepted involved the transfer of the
Zimbabwean population of Nile crocodile to Appendix II in
1983.
Ranching proposals tend to be focused on a particular state,
or operation, and do not constitute mechanisms for the
control of the trade in its entirety. In essence, they continue
the overall controls in effect while allowing very limited
utilisation to recommence under particular conditions.
In 1983, a species-based approach was first adopted with
regard to exploitation of the African Leopard. Although
listed on Appendix I, it was recognised in Conference
Resolution 4.13 that specimens of the leopard could be
killed "to enhance the survival of the species". With this,
the Conference of the Parties approved an annual quota of
460 specimens, and allocated these between the range
states. In 1985 this quota was then increased to 1,140
animals, and in 1992 to 2,055.
This approach to trade management was extended in 1985
with Resolution 5.21, which provided for the systematic
transfer from Appendix I to Appendix II of populations
where the countries of origin agree a quota system which is
sufficiently safe so as to not endanger the species. Five
different species have been subject to quota systems under
this Resolution: three African crocodiles, one Asian
crocodile, and the Asian Bonytongue (a fish) for which
Indonesia was allowed a quota of 1,250 specimens.
None of these trade control systems went further than the
development of species-based quotas. In particular, no
external control structure was ever implemented, this being
left to the discretion of producer states. Thus, predictably,
the quotas can be abused: for example, Indonesia is
believed to have issued permits for about 140% of its first
year’s quota of Bonytongues (Anon., 1991).
The African Elephant management quota system
The third avenue of innovation under CITES, and the most
concentrated attempt thus far to develop an international
control structure within the system, was the creation under
Resolution 5.12 of a Management Quota System for the
African Elephant. This system was founded upon the ideas
of controls based on management decisions taken by
producer countries but enforced by consumer countries.
Annual quotas were to be constructed at the outset of each
year, and producer states were then to issue permits not
exceeding these quotas. Then consumer states were to
disallow all imports unless accompanied by a Management
Quota System permit.
This did not result in an effective control system for one
very important reason. The Management Quota System
provided no external checks on the discretion of the
producer states in determining annual quotas. There were
no externally enforced incentives for sustainable use. This
resulted in most states basing their annual ‘management
quotas’ of ivory on the ‘expected’ confiscations from
poachers. In addition, there were no disincentives for cross-
border exploitation, since consumer states were allowed to
import ivory unquestioningly from any exporter issuing
permits.
The Management Quota System failed as a consequence of
these clear inadequacies, resulting in a collapse of public
International Policy and Legal Assistance
confidence in the capacity for trade controls to work. This
in turn resulted in the transfer of the African Elephant to
Appendix I, despite the fact that there remain approximately
600,000 elephants. Each of these requires about 0.5km? of
grazing lands and in a land where human populations are
doubling every 20 years, it is difficult to maintain existing
diverse resources, especially when their values are reduced
(Barbier‘et al., 1990).
Despite the difficulties experienced by CITES in achieving
its aim of limiting the over-exploitation of wildlife by
international trade, the convention itself has proved very
popular and, with 113 signatories, it is, together with the
World Heritage Convention and Ramsar, among the most
significant examples of international action to preserve
biodiversity.
Future trends
The 20 years in which CITES has been in existence have
indicated the enormous potential that a truly effective
regime could have in the effort to secure the long-term
survival of significant amounts of the world’s biodiversity.
The importance of properly distributing the costs and
benefits of this biodiversity is becoming increasingly
apparent. The effort of ITTO and FAO to move world
tropical timber production on to a sustainable basis is just
one of many examples in which international institutions are
attempting to correct previous distortions in the distribution
of these costs and benefits.
REGIONAL SEAS PROGRAMME
A primary example of what can be achieved by means of
international coordination of national efforts to conserve
biodiversity is the UNEP Regional Seas Programme. The
object of this programme, initiated in 1974, has been to
develop an integrated and comprehensive approach to
protect the marine environment.
Such an approach is necessary because of the nature of
threats to the marine environment. Dumping from ships,
land-based pollution and overfishing are among the threats
which national governments acting unilaterally find difficult
Table 31.4 UNEP Regional Seas Programme: Areas and Action Plans
REGIONAL SEA AREA
ACTION PLAN ADOPTED
PUBLISHED IN RSRS*
Mediterranean February 1975 No. 34 (1983, rev. 1985)
Gulf April 1978 No. 35 (1983)
West/Central Africa March 1981 No. 27 (1983)
Southeast Pacific November 1981 No. 20 (1983)
Red Sea February 1982 No. 81 (1986)
Caribbean April 1981 No. 26 (1983)
Eastern Africa June 1985 No. 61 (1985)
South Pacific March 1982 No. 29 (1983)
East Asia October 1981 No. 24 (1983)
South Asia in preparation
Note: * UNEP Regional Seas Reports and Studies.
501
3. Conservation and Management of Biodiversity
to control. UNEP has sought to develop the necessary
international cooperation through its Regional Seas
Programme, which currently covers 10 different regions,
consists of 24 separate international agreements and
involves over 50 different countries. In a period of less than
20 years the programme has made a major impact on the
conservation of the marine environment. The regions
covered are illustrated in Fig. 31.2, and details of the legal
instruments and the action plans developed for each are
given in Tables 31.4 and 31.5.
The Mediterranean was the first region in which the
programme developed a cooperative framework for
environmental protection. The approach developed here has
served as a blueprint for other regional plans subsequently
developed by UNEP.
The first stage in this process was the development of a
regional action plan. The Mediterranean Action Plan was a
comprehensive interdisciplinary attempt to develop and
implement substantive programmes for the protection of the
marine environment. The Action Plan formed the basis of
the Convention for the Protection of the Mediterranean Sea
Against Pollution, otherwise known as the Barcelona
Convention. This convention has four basic components:
e Environmental assessment
monitoring network
Environmental management through the ‘Blue Plan’ for
coordinated development of the coastal regions and
‘Priority Action Programmes’ for cooperation in coastal
settlements, agriculture, freshwater resources, soils
renewable energy and tourism
Institutional arrangements (such as the establishment of
a permanent secretariat and the regular holding of
conferences of the parties)
Financial arrangementsto help countries implement some
requirements of the convention.
through the MEDPOL
Success of this regional endeavour and of every subsequent
regime developed under the auspices of this programme is
entirely dependent upon the involvement of the majority of
the coastal countries in the regions concerned. In order to
achieve this, the Barcelona Convention was designed to be
as flexible as possible. As a result of this need for
flexibility the convention which the parties adopted was a
framework convention, which outlines in broad terms what
obligations the parties are willing to undertake. These basic
principles are then developed into specific obligations
through the adoption of protocols to the main convention.
Another feature which provides considerable flexibility for
the regulatory regime established pursuant to these
programmes is the use of technical annexes, including
502
‘black lists’ and ‘grey lists’ for substances identified as
potentially harmful to the environment. These lists may be
amended through an accelerated procedure not requiring
diplomatic ratification. This approach has ensured that for
each region the programme has been able to enlist most if
not all of the relevant coastal states.
The Barcelona Convention contains many mechanisms to
foster the active cooperation of all of the contracting
parties. A requirement for periodic conferences of the
parties helps to retain the parties’ interest and keep the
convention from becoming a ‘sleeping treaty’. The
establishment of an active secretariat ensures that there is
continuity in management. The secretariat runs numerous
programmes which are designed to provide support to
parties in implementing the provisions of the convention,
such as: the provision of technical assistance; training
programmes; financial aid; and provision of administrative
support at the periodic conferences. An active
administration has also been important in the dissemination
of new techniques and technology amongst the contracting
parties and from region to region. All of these supporting
measures help develop cooperation between the contracting
parties.
A measure of the success of this programme in developing
cooperation and in protecting the marine environment of
many of the threatened regions can be gained from
comparing its development and that of the UN Convention
on the Law of the Sea (UNCLOS). Both initiatives are
concerned with the marine environment, both were initiated
at the same time and both are international, involving a
wide range of countries. The Regional Seas Programme has
put in place regimes which have already had an impact on
problems in the marine environment, whereas UNCLOS has
yet to enter into force.
References
Anon. (TRAFFIC Japan) 1991. Asian Bonytongue exports from
Indonesia. TRAFFIC Bulletin 12(1,2):3.
Barbier, E.B., Burgess, J.C., Swanson, T.M. and Pearce, D.W. 1990.
Elephants, Economics and Ivory. Earthscan, London.
Cornish, W.R. 1989. Intellectual Property: patents, copyright, trade
marks and allied rights. Sweet and Maxwell, London.
Congress of the United States, Office of Technology Assessment 1990.
New Developments in Biotechnology: patenting life. Marcel Decker,
Inc., New York.
Chapter planned by Timothy M. Swanson. Authors as
follows: Funding, Sam Johnston; Regulated trading in
wildlife products, Shirra Freedman; Intellectual property
rights for biotechnology, Nigel Howard; International
Obligations and Regional Seas programme, Sam Johnston.
International Policy and Legal Assistance
Figure 31.2 UNEP Regional Seas areas
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503
3. Conservation and Management of Biodiversity
Table 31.5 Regional Seas conventions
MEDITERRANEAN
BARCELONA DUMPING EMERGENCY LAND-—BASED PROTECTED AREAS
CONVENTION PROTOCOL? PROTOCOL? SOURCES PROTOCOL"! PROTOCOL
SIGNED/ IN FORCE SIGNED/ IN FORCE SIGNED/ IN FORCE SIGNED/ IN FORCE SIGNED/ IN FORCE
ACCEDED ACCEDED ACCEDED ACCEDED ACCEDED
Algeria 16/03/81 15/04/81 16/03/81 15/04/81 16/02/81 15/04/81 02/05/83 17/06/83 16/05/85 23/03/86
Cyprus 16/02/76 19/12/79 16/02/76 19/12/79 16/02/76 19/12/79 17/05/80 27/12/87 28/06/88 28/06/88
Eygpt 16/02/76 23/09/78 16/02/76 23/09/78 16/02/76 23/09/78 18/05/83 17/06/83 16/02/83 23/03/86
France 16/02/76 10/04/78 * 16/02/76 10/04/78 * 16/02/76 10/04/78 * 17/05/80 17/06/83 * 03/04/82 02/10/86 *
Greece 16/02/76 02/02/79 16/02/76 02/02/79 16/02/76 02/02/79 17/05/80 25/02/87 03/04/82 25/02/87
Israel 16/02/76 02/04/78 * 16/02/76 31/03/84 16/02/76 02/04/78 18/05/80 04/04/82 27/11/87
Italy 16/02/76 05/03/79 16/02/76 05/03/79 16/02/76 05/03/79 17/05/80 03/08/85 03/04/82 23/03/86
Lebanon 16/02/76 12/02/78 16/02/76 12/02/78 16/02/76 12/02/78 17/05/80 n/a n/a
Libya 31/01/77 02/03/79 31/01/77 02/03/79 31/01/77 02/03/79 17/05/80 06/06/89 06/06/89
Malta 16/02/76 12/02/78 16/02/76 12/02/78 16/02/76 12/02/78 17/05/80 03/04/82 10/02/88
Monaco 16/02/76 12/02/78 16/02/76 12/02/78 16/02/76 12/02/78 17/05/80 17/06/83 03/04/82 29/05/89
Morocco 16/02/76 15/02/80 16/02/76 15/02/80 16/02/76 15/02/80 17/05/80 11/03/87 03/04/82 22/06/90
Spain 16/02/76 12/02/78 16/02/76 12/02/78 16/02/76 12/02/78 17/05/80 05/07/84 03/04/82 21/01/88
Syria 26/12/78 25/01/79 * 26/12/78 25/01/79 26/12/78 25/01/79 n/a n/a
Tunisia 25/05/76 12/02/78 25/05/76 12/02/78 25/05/76 12/02/78 17/05/80 17/06/83 03/04/82 23/03/86 *
Turkey 16/02/76 06/05/81 16/02/76 06/05/81 16/02/76 06/05/81 21/02/83 17/06/83 06/11/86 06/12/86
Yugoslavia 15/09/76 12/02/78 15/09/76 12/02/78 15/09/76 12/02/78 30/03/83 23/03/86
EEC 13/09/76 15/04/78 13/09/76 15/04/78 13/09/76 11/09/81 17/05/80 06/11/83 30/03/83 23/03/86
1 Convention for the Protection of the Mediterranean Sea Against Pollution, adopted at Barcelona on 16 February 1976.
2 Protocol for the Prevention of Pollution of the Mediterranean Sea by Dumping from Ships and Aircraft, adopted at Barcelona on the 16 February 1976.
2 Protocol Concerning Cooperation in Combating Pollution of the Mediterranean Sea by Oil and Other Harmful Substances in Cases of Emergency,
adopted at Barcelona on 16 February 1976.
"1 Protocol for the Protection of the Mediterranean Sea Against Pollution from Land—based Sources, adopted on 17 May 1980.
*2 Protocol Concerning Mediterranean Specially Protected Areas, adopted at Geneva on 3 April 1982.
* with reservation.
GULF
KUWAIT EMERGENCY
CONVENTION! PROTOCOL?
SIGNED IN FORCE SIGNED IN FORCE
Bahrain 24/04/78 01/07/79 24/04/78 01/07/79
Iran 24/04/78 01/06/80 24/04/78 ; 01/06/80
lraq 24/04/78 01/07/79 24/04/78 01/07/79
Kuwait 24/04/78 01/07/79 24/04/78 01/07/79
Oman 24/04/78 01/07/79 24/04/78 01/07/79
Qatar 24/04/78 01/07/79 24/04/78 01/07/79
Saudi Arabia 24/04/78 26/03/82 24/04/78 26/03/82
United Arab Emirates 24/04/78 01/03/80 24/04/78 01/03/80
! Kuwait Regional Convention for Co—operation on the Protection of the Marine Environment from Pollution, adopted on 23 April 1978.
2 Protocol Concerning Regional Co—operation in Combating Pollution by Oil and Other Harmful Substances in Cases of Emergency, adopted 23 April 1978.
WEST/CENTRAL AFRICA
ABIDJAN EMERGENCY
CONVENTION' PROTOCOL?
SIGNED/ IN FORCE SIGNED/ IN FORCE
ACCEDED ACCEDED
Benin 23/03/81 23/03/81
Cameroon 01/03/83 05/08/84 01/03/83 05/08/84
Congo 23/03/81 23/03/81
Cote d'ivoire 23/03/81 05/08/84 23/03/81 05/08/84
Gabon 23/03/81 23/03/81
Gambia 23/03/81 05/02/85 23/03/81 05/02/85
Ghana 23/03/81 23/03/81
Guinea 23/03/81 05/08/84 23/03/81 05/08/84
Liberia 23/03/81 23/03/81
Mauritania 22/06/81 22/06/81
Nigeria 23/05/81 05/08/84 23/05/81 05/08/84
Senegal 23/03/81 05/08/84 23/03/81 05/08/84
Togo 23/03/81 05/08/84 23/03/81 05/08/84
! Convention for Co—operation in the Protection and Development of the Marine and Coastal Environment of the West and Central African Region,
adopted at Abidjan on 23 March 1981.
Z Froieral Concerning Co-operation in Combating Pollution in Cases of Emergency in the West and Central African Region, adopted at Abidjan on 23
March 1981.
SOUTH-EAST PACIFIC
LIMA EMERGENCY SUPPLEMENTATY LAND-—BASED
CONVENTION' AGREEMENT? PROTOCOL? SOURCES PROTOCOL
SIGNED IN FORCE SIGNED IN FORCE SIGNED IN FORCE SIGNED IN FORCE
Chile 12/11/81 19/05/86 12/11/81 14/07/86 22/07/83 20/05/87 22/07/83 23/09/86
Colombia 12/11/81 19/05/86 12/11/81 14/07/86 22/07/83 20/05/87 22/07/83 23/09/86
Ecuador 12/11/81 19/05/86 12/11/81 14/07/86 22/07/83 11/01/88 22/07/83 11/01/88
Panama 12/11/81 21/09/86 12/11/81 21/09/86 22/07/83 20/05/87 22/07/83 23/09/86
Peru 12/11/81 25/02/89 12/11/81 22/07/83 22/07/83
' Convention for the Protection of the Marine Environment and Coastal Area of the South—East Pacific, adopted at Lima on 12 November 1981.
? Agreement on Regional Co—operation in Combating Pollution of the South—East Pacific by Hydrocarbons or Other Harmful Substances in cases of
Emergency, adopted at Lima on 12 November 1981.
3 Supplementary Protocol to the Agreement on Regional Co—operation in Combating Pollution of the South—East Pacific by Hydrocarbons or Other
Harmful Substances in Cases of Emergency, Adopted at Quito on 22 July 1983.
*! Protocol for the Protection of the South—East Pacific Against Pollution from Land—Based Sources, adopted at Quito on 22 July 1983.
504
International Policy and Legal Assistance
Table 31.5 Regional Seas conventions (continued)
RED SEA
JEDDAH EMERGENCY
CONVENTION! PROTOCOL2
SIGNED IN FORCE SIGNED IN FORCE
Eygpt 21/05/90 20/08/90
Jordan 14/02/82 07/02/89 14/02/82
Saudi Arabia 14/02/82 20/08/82 14/02/82 20/08/82
Somalia 14/02/82 30/05/88 14/02/82
Sudan 14/02/82 20/08/82 14/02/82 20/08/82
Yemen 14/02/82 20/08/82 14/02/82 20/08/82
1 Regional Convention for the Conservation of the Red Sea and Gulf of Aden Environment, adopted at Jeddah on 14 February 1982.
2 Protocol Concerning Regional Co—operation in Combating Pollution by Oil and Other Harmful Substances in Cases of Emergency, adopted at Jeddah on
14 February 1982.
CARIBBEAN
CARTAGENA SPECIALLY PROTECTED OW SPILLS
CONVENTION AREAS PROTOCOL? PROTOCOL?
SIGNED/ IN FORCE SIGNED/ IN FORCE SIGNED/ IN FORCE
ACCEDED ACCEDED ACCEDED
Antigua and Barbuda 11/09/86 11/10/86 18/01/90 11/09/86 11/10/86
Barbados 05/03/84 11/10/86 18/01/90 05/03/84 11/10/86
Colombia 24/03/83 18/01/90 24/03/83
France 24/03/83 11/10/86 * 18/01/90 24/03/83 11/10/86 *
Grenada 24/03/83 16/09/87 18/01/90 24/03/83 16/09/87
Guatemala 05/07/83 18/01/90 05/07/83
Honduras 24/03/83 18/01/90 24/03/83
Jamaica 24/03/83 01/05/87 18/01/90 24/03/83 01/05/87
Mexico 24/03/83 11/10/86 18/01/90 24/03/83 11/10/86
Netherlands"! 24/03/83 11/10/86 18/01/90 24/03/83 11/10/86
Nicaragua 24/03/83 18/01/90 24/03/83
Panama 24/03/83 06/11/87 24/03/83 06/11/87
St Lucia 24/03/83 11/10/86 18/01/90 24/03/83 11/10/86
Trinidad and Tobago 24/03/83 11/10/86 18/01/90 24/03/83 11/10/86
United Kingdom®2 24/03/83 11/10/86 * 18/01/90 24/03/83 11/10/86 *
United States 24/03/83 11/10/86 18/01/90 24/03/83 11/10/86
Venezuela 24/03/83 17/01/87 18/01/90 24/03/83 17/01/87
EEC 24/03/83 18/01/90
1 Convention for the Protection and Development of the Marine Environment of the Wider Caribbean Region, adopted at Cartagena on 24 March 1983.
2 Protocol Concerning Specially Protected Areas and Wildlife to the Convention for the Protection and Development of the Marine Environment of the
Wider Caribbean Region, adopted in Kingston on 17 January 1990.
3 Protocol Concerning Co-operation in Combating Oil Spills in the Wider Caribbean Region, adopted at Cartagena on 24 March 1983.
™! On behalf of Aruba and the Netherlands Antilles Federation.
*2 On behalf of the Cayman Islands and the Turks and Caicos Islands, reserving the right to include at a future date the other territories of the United
ssiiedom participating in the Caribbean Action Plan (Anguilla, British Virgin Islands and Montserrat).
* with reservation.
EASTERN AFRICA
NAIROBI PROTECTED AREAS EMERGENCY
CONVENTION' PROTOCOL2 PROTOCOL?
SIGNED IN FORCE SIGNED IN FORCE SIGNED IN FORCE
France™! 21/06/85 21/06/85 21/06/85
Kenya 11/09/90 11/09/90 11/09/90
Madagascar 21/06/85 21/06/85 21/06/85
Seychelles 21/06/85 21/06/85 21/06/85
Somalia 21/06/85 21/06/85 21/06/85
EEC 19/06/86 19/06/86 19/06/86
1 Convention for he Protection, Management and Development of the Marine and Coastal Environment of the Eastern African Region, adopted at Nairobi
on 21 June 1985.
2 Protocol Concerning Protected Areas and Wild Fauna and Flora in the Eastern African Region, adopted at Nairobi on 21 June 1985.
a peaiecol Concerning Co-operation in Combating Marine Pollution in Cases of Emergency in the Eastern African Region , adopted at Nairobi on 21 June
1985.
*! On behalf of Réunion.
SOUTH PACIFIC
NOUMEA EMERGENCY DUMPING
CONVENTION' PROTOCOL? PROTOCOL?
SIGNED IN FORCE SIGNED IN FORCE SIGNED IN FORCE
Australia 24/11/87 22/08/90 24/11/87 24/11/87
Cook Islands 25/11/86 22/08/90 25/11/86 25/11/86
Federated States of 09/04/87 22/08/90 09/04/87 09/04/87
Micronesia
France 25/11/86 22/08/90 25/11/86 25/11/86
Marshall Islands 25/11/86 22/08/90 25/11/86 25/11/86
Nauru 15/04/87 15/04/87 15/04/87
New Zealand 25/11/86 22/08/90 25/11/86 25/11/86
Palau 25/11/86 25/11/86 25/11/86
Papua New Guinea 03/11/87 22/08/90 03/11/87 03/11/87
Tuvalu 14/08/87 14/08/87 14/08/87
United Kingdom 16/07/87 16/07/87 16/07/87
United States 25/11/86 10/07/91 25/11/86 25/11/86
Samoa 25/11/86 22/08/90 25/11/86 25/11/86
1 Convention for the Protection of Natural Resources and Environment of the South Pacific Region, adopted at Noumea on 25 November 1986.
2 Protocol Concerning Co-operation in Combating Pollution Emergencies in the South Pacific Region, adopted at Noumea on 25 November 1986.
3 Protocol for the Prevention of Pollution of the South Pacific Region by Dumping, adopted at Noumea on 25 November 1986.
505
3. Conservation of Biodiversity
32. INTERNATIONAL AID
As noted in the previous chapter, national efforts alone are
not sufficient, despite their fundamental importance, to
ensure adequate management of biological diversity.
Chapter 31 surveyed international support for national
measures as provided by policy and legal assistance; this
chapter will discuss the role of direct development aid.
Even though the proportion of total international aid which
is specifically targeted for the conservation of biodiversity
is relatively small, it nonetheless plays a very important
role. The three principal kinds of assistance examined here
are: international development assistance, international
programmes for the conservation of tropical forests, and a
form of debt purchase widely known as ‘debt-for-nature’
exchange.
The impact on biodiversity of international development
assistance, whether intended or not, is felt through many
channels and will vary in nature and extent. The following
review of this international assistance attempts to assess to
what extent bilateral and multilateral funds are directly
targeting biodiversity conservation, and how this is
incorporated into the project and loan appraisal process.
An examination of two international programmes designed
to conserve tropical forests highlight some of the problems
faced by any international effort to conserve biodiversity.
Given the cross-sectoral nature of environmental issues and
the number of different international programmes currently
established, even where there is some consensus over what
action is required, there still remain major problems of
organisation and coordination to be overcome.
The last part of this chapter shows that there is no single
and ready solution to the problem of conserving
biodiversity. Several years ago ‘Debt-for-Nature’ exchanges
were regarded as a major chance to counter many of the
underlying causes of biodiversity degradation. As this
mechanism has developed it has become evident that some
predictions of its importance were over-optimistic.
INTERNATIONAL DEVELOPMENT ASSISTANCE
The term ‘international development assistance’ (IDA)
(otherwise ‘overseas development assistance’ or ODA) is
used here to refer to concessional aid provided by
governments to developing countries. IDA is delivered
directly by donor countries’ bilateral aid or by multilateral
institutions such as the United Nations and the major
development funds and banks. Concessional aid includes
grants and loans made at less than market interest rates, but
not other types of official financial flows such as export
credits, grants by private voluntary agencies or private
flows at market terms. World IDA accounted for roughly
half of the $110 billion in financial resources transferred to
developing countries in 1989. Non-concessionary bilateral
and multilateral disbursements (14%), foreign direct
investment (20%) and international bank lending (7%) make
up large portions of the remaining resources transfers.
Resource transfers to the developing countries are only
approximately half as large in real terms as they were at
506
their peak in 1981. At that time 38% of resource transfers
consisted of private international bank lending. The drain on
development finance resulting from the ensuing debt crisis
of the 1980s is far from over. In 1989 developing countries
paid out interest and dividends of $108 billion - a figure
roughly equal to the incoming financial resources cited
above. Total overseas development assistance over this
period has remained fairly stable. While multilateral
disbursements have been unchanging, Arab donors’
contributions have fallen dramatically from close to $10
million in 1980 to less than $2 million in 1989. The group
of 18 countries making up the Development Assistance
Committee (DAC) of the Organization for Economic
Cooperation and Development (OECD) have made up for
this shortfall by steadily increasing their IDA expenditures.
Commitments by DAC countries (listed in Table 32.1) have
risen from an average of 62% of world IDA over the period
1980-81 to 87% in 1988-89.
The DAC average of contributing 0.35% of GNP to IDA
masks considerable discrepancy between the performance of
the USA and Japan, on the one hand, and Scandinavian and
Dutch donors on the other. Table 32.1 clearly reveals that
the average Norwegian contributes more than six times as
much IDA as does the average American. The official DAC
target of 0.7% of GNP, first proposed by the Pearson
Commission in 1969 remains elusive for the majority of
DAC members. Nonetheless, official flows of IDA remain
the primary vehicle of aid transfers. In comparison, the
average for aid from non-governmental organisations in
1988-89 was estimated at just 0.03% of GNP.
Two other important IDA statistics measuring aid
performance are the ‘grant element’ and the status of ‘tied
aid’. In recent years only a couple of DAC countries failed
to achieve the DAC target of delivering 86% of their funds
on grant terms. Tying aid to purchases of goods or services
from the donor country remains a more complicated and
controversial topic. Tied aid benefits donors while inflating
the cost of recipient country purchases by 10% to 20% (de
Silva, 1982). The disparity between tied aid figures in
Table 32.1 reveals the lack of coherent progress within the
DAC on this issue. In 1988 approximately 30% of DAC
country commitments to IDA were tied and another 7.5%
partially untied.
The impact of international development assistance on
biodiversity is felt through many channels: projects, sectoral
and macroeconomic policy reform, and institutional and
human resource initiatives. The list is long and the impacts
on biodiversity, intentional and otherwise, will vary in
direction and magnitude.
The following review of bilateral and multilateral aid
policies on biodiversity bypasses specific negative impacts
of IDA on biodiversity, which are well documented
elsewhere, in favour of finding evidence of bilateral and
multilateral action towards the conservation of biodiversity.
A relatively new concern, such as biodiversity, is likely to
be incorporated into the development assistance process
either through increased funding for projects, technical
cooperation and other means of addressing the issue, or by
International Aid
Table 32.1 International development assistance
COUNTRY NET AMOUNT SHARE OF SHARE OF TIED AID AS
AMOUNT PER CAPITA GNP WORLD IDA % OF TOTAL
1988 1988 1988-89 mean 1988-89 mean IDA?
(US$million) (US$) % %
Ireland 53.7 1522 0.2 0.1 (3.8)
New Zealand 96.6 29.4 0.2 0.2 36.4
USA 8,749.2 35.3 0.2 15.9 37.7
Austria 298.5 39.2 0.2 0.5 68.8
UK 2,640.2 46.2 0.3 4.8 46.4
Italy 3,383.9 58.9 0.4 6.2 57.6
Australia 1,018.4 61.1 0.4 1.9 32.7
Belgium 662.9 67.0 0.4 er (34.1)
Japan 9,312.5 75.8 0.3 16.9 11.5
Germany 4,946.9 80.1 0.4 9.0 32.5
Canada 2,238.3 85.8 0.5 4.1 34.5
Switzerland 609.8 91.0 0.3 1.1 15.0
Finland 643.3 129.9 0.6 1.2 47.1
France! 7,289.9 130.1 0.8 13.3 37.7
Netherlands 2,229.4 150.6 1.0 4.1 10.4
Denmark 950.1 185.2 0.9 137. 15.8
Sweden 1,645.3 195.0 0.9 3.0 21.2
Norway 970.1 229.9 1.1 1.8 20.8
Total DAC 47,739.0 66.7 0.3 86.6 (30.5)
Total Non-DAC OECD 382.0 - 0.1 0.7 -
Total Arab Donors 1,900.0 - 0.7 3.5 -
Central and Eastern Europe 4,534.0 - - 8.2 -
Total non-Arab LDC donors 425.0 - 0.0 0.8? -
WORLD TOTAL 54,980.0 - - 100.0 -
Source: DAC 1990. Development Co-operation: efforts and policies of the members of the Development Assistance Committee. OECD, Paris.
Notes: () are DAC estimates. ' Including aid to French possessions. * Figures for 1988 only.
inclusion in assessment procedures for projects that may
have unintended or indirect impacts on biodiversity. For
this reason the emphasis below is on reviewing whether or
not, and how, biodiversity is being incorporated into the
project/loan appraisal process and whether funds are
targeting biodiversity directly.
Unfortunately, biodiversity issues crop up in a number of
traditional sectors, such as health, energy, agriculture,
mining, transport, food aid, etc., making it difficult to
clearly define a biodiversity ‘sector’ or to sum up
biodiversity expenditures. This obstacle is one shared by
environmental issues in general - indeed a further confusion
may occur between environmental issues and biodiversity
issues broadly defined. For this reason, the approach taken
below is to report on progress specific to biodiversity where
such information is available. An explicit role for
biodiversity in project and policy formulation, or in funding
commitments, may be one indicator of the level of
awareness and seriousness of commitment in donor agencies
towards conserving biodiversity. The intention is not to
recommend that biodiversity merits it own ‘sectoral’ billing,
but to begin the process of evaluating the resources that
development agencies are committing to conservation.
When specific actions on biodiversity are not evident,
progress on the general environmental front is documented
as indicative of growing concern that may soon encompass
biodiversity directly.
507
BILATERAL DEVELOPMENT ASSISTANCE
Funding
Establishing exactly what constitutes funding for
biodiversity is not straightforward. Abramovitz (1991)
compared the results of two studies on US-based funding
for biodiversity conservation carried out by the World
Resources Institute’s Center for International Development
and Environment. Aggregating project-level data from
government sources, NGOs, foundations, universities, etc.,
the total investment came to $37.5 million for 1987 and
$62.9 million for 1989. Listed below are the major
categories of activity areas used in classifying all 1,093
projects recording for 1989:
Research Policy Planning and Analysis
Basic Conservation Law/Regulatory
Ecosystem Natural Resources Accounting
Species Economic Policy Analysis
Applied Program/Project Planning
Response to Disturbance Program/Project Design
Statistics, Indicators
Environmental Impact Assessment
Education
Public Awareness
Curriculum Development
Degree Training
Technical Training
Institutional Support
Economic Assessment
Social/Cultural Values
Systematics/Inventory
Site or Species Management
Protected Areas
Planning and Assessment
Management
Buffer Zone Management
Ecosystem Restoration
Species Management - in situ, ex situ
3. Conservation of Biodiversity
While this study is the most comprehensive effort to date,
even such a detailed survey cannot avoid the difficulties
inherent in extracting biodiversity project data from
programmes aligned along traditional sectoral or country
boundaries. In addition, the non-equivalence of biodiversity
and biological resources means that some activities will be
slotted in as biodiversity funding when they have little to do
with diversity per se. Determining whether or not a project
is concerned with management of biological resources
generally, or more specifically with biodiversity, is not an
easy task. The difficulty with relaxing the emphasis on
diversity is that the difference between funding for
biodiversity and funding for the environment will become
more and more blurred.
It is more difficult to categorise the importance with regard
to funding levels of projects that have only a secondary or
tertiary focus on biodiversity. Often, the perception that
biodiversity pertains only to wild species, ex situ gene
storage or protected areas (as in the WRI study) may also
divert attention from the role of species or genetic diversity
in production systems. Agriculture, aquaculture, forestry,
fishery and other rural development projects may have a
significant diversity component. On the other hand, projects
aimed at developing sustainable resource management may
involve some loss of biodiversity.
Recognising the inexactness of even the most
comprehensive study, the London Environmental Economics
Centre (LEEC) conducted a brief survey of DAC bilateral
agencies to obtain a general impression of the visibility of
biodiversity as a concern in agency funding and project
appraisal. None of the agencies responding to the
questionnaire currently disaggregate their expenditures to
identify the amount spent on biodiversity or genetic
resources. As shown in Table 32.2 roughly half of the
agencies do calculate the extent of funding for the
environment. Three others indicated that they will be doing
so in the near future. The movement towards identification
of environmental expenditures is a natural precursor to
establishing similar reporting procedures for biodiversity.
Another way in which awareness regarding biodiversity
may be manifested is through setting aside funds for use
specifically on biodiversity. Since 1983 the US Congress
has earmarked funds for biodiversity conservation in
USAID’s annual appropriation. Recently, a number of DAC
agencies are now earmarking funds for biological diversity
as described in Table 32.2.
The difficulty in interpreting such aggregate numbers is
illustrated in the case of the data provided by Germany’s
BMZ (see Table 32.2 notes). While BMZ has earmarked
DM3.5 million for biodiversity in 1991 they also reported
Table 32,2 Bilateral funding for biodiversity and the environment
DONOR BILATERAL BIODIVERSITY ARE EXPENDITURES ON THE
COUNTRY FUNDS - 1988 FUNDS SET ASIDE ENVIRONMENT IDENTIFIED?
$US million $US million $US million
Australia’ 622 AS 4.3m in 1991/92 A$ 10.6m in 1990/91 9
Austria 162 none - yes, not available -
Belgium 415 none - will identify soon =
Canada 1,583 none - no =
Denmark 478 US$ 0.115m in 1991 (0) no -
DKr 25m in 1992-96 4
France 5,601 ? plan to - FF 200m in 1991 35
Germany (BMZ) 3,172 ° DM 3.5m in 1991 2 DM 1020m in 1990* 610
(GTZ) none - DM 200m in 1990 120
lreland 22 none - will identify soon -
Italy 2,408 none - will identify by 1992 -
Netherlands ioe e - US$ 135m in 1991 135
New Zealand 93 yes, not assessed - NZ$ 50m in 1990/91 28
Norway 570 none = NOK 852 in 1990° 130
Sweden 1,034 SKr 1.5m in 1991/92 {e) no -
Switzerland 4147 - no -
UK 1,430 8 - no® -
usa’? 6,765 US$ 9.9m in 1989 9 no -
Source: LEEC questionnaire to DAC members. Responses received from all but Finland, Germany - KfW and Japan. EEC responses are presented
under the section on multilaterals. Exchange rates used are from the rates listed in the Financial Times on 30 September 1991
Notes: ' Directly targeted funds only. ? Including aid to French possessions. * Total for all three German agencies: Bundesminister flir wirtschaftliche
Zusammenarbeit (BMZ), Deutsche Gesellschaft fir Technische Zusammenarbeit (GTZ) and Kreditanstalt fiir Wiederaufbau (KfW). * Commitments
not expenditures; does not include DM325m on tropical forests. * No funds are set aside, but expenditures on biodiversity in 1991 came to roughly
$US 5m. * Includes NOK 651m on environmental integrated projects and NOK 201m on direct environmental projects. Excludes NOK 315m on
population.’ Although expenditures are not specified, the conservation of biodiversity constitutes a basic objective of the Directorate’s development
policy. * Although ODA does not set aside funds it does conduct strategic work on biodiversity conservation and finances many biodiversity projects;
for example, ODA is the primary funder of this report. ? ODA prefers to integrate environmental issues throughout its aid programme rather than
treat it as a separate sector and keep separate statistics on it. '° The figure for USAID represents actual expenditures as reported by the Biodiversity
Projects Database maintained by the World Resources Institute.
508
forest sector expenditures of DM325 million in 1990.
Setting aside money explicitly conveys BMZ’s concern for
biological diversity. However, the size of BMZ’s existing
efforts in tropical forestry indicate that the impact of the set
aside funds is likely to be superseded by the funds allocated
for the ‘preservation and development’ of tropical forests.
The positive impact of the amount set aside will be small
relative to the forestry expenditures if a reasonable
percentage of the forestry funds goes towards preservation
or sustainable use of the forests. On the other hand, if a
large portion of forest sector expenditures support
unsustainable logging activities, the overall negative impacts
of these expenditures would overwhelm the potential
benefits gained from the earmarked funds. Clearly the direct
impact of donor allocations for biodiversity shown in Table
32.2 (all of them less than $10 million) will be negligible
compared to the real effects of much larger sectoral
expenditures on forests, agriculture, transport, etc. The
significance of the allocation would be increased by using
the money to sponsor innovative projects, research or
institutional initiatives.
Project appraisal
Ideally, project appraisal procedures should act as a control
on sectoral projects with significant impacts on biodiversity.
If projects are screened for negative impacts on biodiversity
then funding allocated towards biodiversity conservation
will assume greater importance instead of generating
suspicion that they are just compensation for the ill-effects
of the remaining development portfolio. Table 32.3 reveals
that most DAC agencies already undertake environmental
impact assessments of project and loan proposals.
Increasingly, countries are following the lead of Canada and
the USA in involving host country officials and experts in
the evaluation procedure.
International Aid
Table 32.3 also lists a number of countries that include
evaluation of project impacts on biodiversity into the
appraisal process. Efforts to carry the physical data through
to a complete cost-benefit analysis by monetising the
impacts on the environment and biodiversity is limited.
Both BMZ and the UK’s Overseas Development
Administration (ODA) report that assessment of the
economic effects is undertaken in particular situations.
Given that empirical work detailing the economic value of
biodiversity, and environmental resources in general, is still
an area of front-line research, it is unlikely that full cost
benefit analysis is likely for each and every development
project in the foreseeable future.
MULTILATERAL DEVELOPMENT ASSISTANCE
Multilateral disbursements of overseas development
assistance and non-concessional finance come primarily
from the World Bank Group, the regional development
banks and the UN specialised agencies. Table 32.4 presents
data on the net disbursements by these organisations. Most
of the multilaterals belong to the Committee of International
Development Institutions on the Environment (CIDIE)
which coordinates multilateral activities on the environment.
This section will review the specific achievements on the
environment and biodiversity as reported by CIDIE
members, and then look at the newest and potentially
largest source of multilateral funding for biodiversity
conservation: the Global Environmental Facility.
Heavily criticised for supporting, amongst other activities,
large hydroelectric projects and logging schemes, the World
Bank has gradually developed a policy on environmental
assessment. The 1989 Environmental Assessment
Operational Directive (EAOD) formalises Bank policy on
environmental impact assessment (EIA). The EAOD
Table 32.3 Bilateral agency loan and project appraisal policies
COUNTRY APPRAISAL OF IMPACTS IN PHYSICAL OR MONETARY TERMS
ON THE ENVIRONMENT ON BIODIVERSITY
Australia physical’ no
Austria physical no
Belgium soon no
Canada physical physical?
Denmark physical physical - soon
France physical no
Germany (BMZ)° physical and monetary physical and monetary
Germany - GTZ physical no
Ireland physical no
Italy physical no
Netherlands physical physical
New Zealand physical physical
Norway ‘physical no
Sweden physical physical
Switzerland physical physical
UK* physical and monetary physical and monetary
Source: LEEC Questionnaire on Biodiversity to DAC members.
Notes: ' With some attempts at monetary evaluation.” Impacts on wildlife flora/fauna and its habitat. * In physical or monetary terms ‘if applicable’.
* In physical and ‘where possible’ in monetary terms.
3. Conservation of Biodiversity
Table 32.4 Multilateral
assistance
development
NET DISBURSEMENTS IN 1988
$US million
CONCESSIONAL NON-CONCESSIONAL
Major Financial Institutions
World Bank Group
IBRD - 3,417
IDA 3,567 --
IFC - 356
Regional Banks/Funds
CEC (or EEC) 2,587 56
IDB 134 1,093
Asian 707 598
African 351 625
IFAD 102 -
Other 65 374
United Nations
UNDP 914 -
WFP 878 -
UNEP’ (59)
Other UN 1,984 -
TOTALS 11,348 6,519
Sources: DAC 1990. Development Co-operation: efforts and policies
of the members of the Development Assistance Committee. OECD,
Paris; UNEP 1990. UNEP Profile. UNEP, Nairobi.
Notes: ' 1989 figure showing funds sourced from regular UN budget
(10%), trust funds (15%), counterpart contributions (8%) and
voluntary contributions to the Environment Fund (67%). IBRD =
International Bank for Reconstruction and Development; IDA =
International Development Administration; IFC = _ International
Finance Corporation, CEC = Commission of the Economic
Community; IDB = Inter-American Development Bank; IFAD =
International Fund for Agricultural Development; UNDP = United
Nations Development Programme; WFP = World Food Programme.
separates projects by type into four categories each of which
receives varying degrees of assessment (Table 32.5).
Environmental projects are generally exempt from EIAs. In
order to determine the nature of the EJA required, if any,
World Bank projects (or components) are classified
according to the likely environmental impacts of the project.
EIA (or EA in the Bank’s terminology) is considered a
flexible procedure that is responsive to the individual
project. The assessment may cover not only environmental
impacts but those on health, cultural property, tribal people
and the environmental impact of resettlement. EIAs should
cover existing conditions, potential direct and indirect
impacts, comparison of project with alternatives,
compensatory measures, environmental management and
training, and monitoring. Wherever possible the costs and
benefits of these elements should be quantified.
As with the economic, financial, institutional and
engineering analyses, the EIA is the borrower’s
responsibility. The Bank provides assistance in designing
the terms of reference for the EIA and normally a field visit
by Bank staff is suggested. The Bank usually recommends
that borrowers hire experts not involved in the project to
carry out the EIA. EIAs for large projects may take up to
18 months to be completed with input from the ongoing
EIA occurring at relevant points in the overall project cycle.
The final report is submitted to the Bank for consideration
with the project or loan application. Funding for ELAs may
be accomplished by a Bank loan or grant and usually comes
to 5-10% of the cost of project preparation.
In its Forest Policy Paper adopted on 18 July 1991 the Bank
has stated that it will not fund commercial logging in
Table 32.5 World Bank project categories and EIA requirements
CATEGORY A: Projects with diverse and significant impacts. EIA required unless directed towards rehabilitation, improved
operation and maintenance, and limited upgrading of facilities.
Aquaculture/Mariculture (LS)
Dams and Reservoirs
Electrical Transmission (LS)
Forestry
Industrial Plants (LS) and Estates
Irrigation and Drainage (LS)
Land Clearance and Levelling
Mineral Development
Pipelines (oil, gas and water)
Port and Harbour Development
Reclamation and New Land
Resettlement
River Basin Development
Rural Roads
Thermal and Hydropower
Tourism (LS)
Transportation infrastructure
Urban Development (LS)
Urban Water Supply/Sanitation (LS)
Manufacture, Transportation and
Projects with serious accident risks
CATEGORY B: Projects which may have specific impacts. Limited EIA required.
Agroindustries (SS)
Aquaculture/Mariculture (SS)
Electrical Transmission (SS)
Industries (SS)
Irrigation and Drainage (SS)
Mini Hydro-Power
Public Facilities
Renewable Energy
Rural Electrification
Telecommunications
Tourism (SS)
Urban Development (SS)
Rural Water Supply/Sanitation
CATEGORY C: Projects which normally do not have serious impacts. ElA normally unnecessary.
Health
Nutrition
Education
Family Planning
Institutional Development
Technical Assistance
CATEGORY D: Projects with a major environmental focus. EIA normally unnecessary
Source: The World Bank, 1989. Operational Directive 4.00, Annex A: Environmental Assessment.
Notes: LS = large-scale projects. SS = small-scale projects.
tropical moist forests. Full EIAs will be required for all
infrastructure projects that may affect tropical moist forests
or other primary forest. This is nothing new since the
EAOD already lists roads, dams and mines as requiring
EIAs. In the 1990 fiscal year the Bank reports that 11 free-
standing environmental projects and 107 loans with
environmental components were approved. The Bank’s
influence on conservation issues will be felt most directly
through its major role in the operation of the new Global
Environmental Facility and involvement in the revised
Tropical Forestry Action Plan.
The Commission of the European Community (CEC) is
integrating environmental considerations into their appraisal
process for projects and programmes. In the future the CEC
plans on incorporating into its evaluations not just physical
EIA but also monetary estimation of environmental impacts.
The European Development Fund (EDF) is financing $26
million worth of environmental training programmes in
West Africa. The Commission does not provide its own
staff with training in the environment, preferring to hire
staff with the required qualifications.
Currently, the CEC does not identify its expenditures on the
environment, nor are there funds specifically for
biodiversity conservation. The Commission has an extensive
research agenda which includes the conservation of
biodiversity and tropical forests, and marine and freshwater
ecology as priority themes.
In 1991 the Commission had roughly $14 million and $2.5
million for work on ‘ecology and developing countries’ and
‘tropical forests’ respectively. The Commission has a $60
million small grants facility at its disposal which frequently
funds grassroots natural resource management initiatives
and encourages cooperation with developed and developing
country NGOs.
Over the period 1990-1995, the CEC will commit $14
billion to activities in African, Caribbean and Pacific
countries. The vast majority of this assistance will be
channelled through the Lome IV Convention mechanism
under which international assistance is provided by the
stabilisation and support of commodity prices for the raw
materials of the less developed parties to the convention. Up
to 75% of the programming undertaken to date has
identified the environment as a key sector. A doubling of
Asian and Latin American funding was accompanied by
allocating $300 million over the next five years (10% of
total funds) to environmental programmes.
The Inter-American Development Bank (IDB) has both a
managerial-level Environmental Committee (CMA) and an
Environmental Protection Division. The latter was created
in 1990 under the Project Analysis Department in order to
ensure that IDB operations comply with member country
legislation and the guidelines on EJA developed by the
Bank. EIAs are required for the proportion (about three-
quarters) of IDB projects that have minor or major
environmental impacts. In 1990 the costs of undertaking
such project reviews were estimated to require an extra $75
million over the next three years. The Bank is also
interested in increasing funding for technical assistance and
NGOs, and promoting debt swaps for environmental
protection.
S11
International Aid
Over the past few years the Asian Development Bank
(AsDB) has upgraded its Environment Unit first to the
status of a Division and most recently into the Office of the
Environment. Accompanying this upgrading of status the
AsDB has added five professional staff and approved a five-
year programme to upgrade environmental awareness and
skills among Bank staff. Guidelines for incorporating EIAs
into the Bank’s project cycle have been developed. In 1989,
30 loans and 43 technical assistance projects had large
environmental components. The AsDB has sponsored
research into the effect of projects on ecologically sensitive
areas, developed guidelines for assessing the impact of
projects on biodiversity and providing technical assistance
funds to encourage its developing country members to
conserve biodiversity. The AsDB has _ incorporated
biodiversity conservation into its Forest Sector Policy
Paper, but the tension between increasing forest production
and conservation remains.
The African Development Bank (AfDB) established an
Environment Unit in 1987 and in June 1990 the AfDB’s
Environment Policy Paper was approved by the Board of
Directors. The Paper prevides guidelines for environmental
impact assessment of both project and non-project loans,
and includes a brief on biodiversity. Review of the AfDB’s
1988 loans in the transport, public utilities and agriculture
sectors (67% of the Bank’s loans) indicated that half of the
loans would have negative environmental impacts. In 1989
just five full EIAs were conducted by the Environment
Unit. During the 1987-1989 period the AfDB reports that
environmentally beneficial projects and environment-linked
projects more than doubled. In 1989 these projects
accounted for 18% of AfDB commitments.
The traditional focus of the International Fund for
Agricultural Development (IFAD) on rural poverty has
recently been broadened in an effort to integrate the
environmental dimension into the Fund’s work programme.
For this reason IFAD has rejected setting up an
environmental unit and is concentrating on upgrading the
knowledge and skills of existing staff. In 1991 IFAD
initiated a two-year programme to develop and test the
introduction of EJA into the project cycle. Preparation of
guidelines for sustainable agriculture and sectoral studies of
resource management is also under way.
The Nordic Investment Bank (NIB) lends roughly $1 billion
per year, 20% of which goes to non-Nordic countries. In
1989-90 the NIB co-financed the Mauritius Environmental
Master Plan and an afforestation project in Indonesia. Brief
environmental appraisals of all NIB loans are conducted by
project officers. The Nordic Environmental Finance
Corporation (NEFCO) began operations in 1990 and is
administered by the NIB. NEFCO provides financing to
joint ventures in ex-Eastern Bloc countries that provide
products beneficial to the Nordic environment.
The United Nations Development Programme (UNDP)
began developing its Environmental Management Guidelines
in 1989. The Guidelines provide the means for non-
specialists to incorporate the principles of environmental
management into their work. UNDP has increased the
coverage of what it considers as ‘environmental projects’
from those that are purely environmental in a scientific
3. Conservation of Biodiversity
sense to include those that encourage sustainable
development and the improvement of the “quality of human
life’. Under this definition UNDP expenditures on
environmental activity totalled US$600 million in 1990. In
1988 expenditures on projects characterised as biodiversity
conservation projects came to $1.5 million. Adding in
subtotals for plant resources and wildlife management the
total spent on biodiversity increases to US$6.7 million.
UNDP is currently an active partner in the Global
Environmental Facility which is described below.
The World Food Programme (WFP) of the United Nations
commits one-third of its ‘food aid’ finance to environmental
project components. WFP is training its staff in
incorporating environmental concerns into project design
rather than approaching environmental issues by way of
rigorous EIAs. A simple checklist assessment has been
devised to alert staff to potential environmental risks during
the project preparation and planning stages. The checklist
does contain cautions regarding the loss of genetic diversity
as cropping patterns change, but does not include specific
diversity considerations within areas such as afforestation,
road construction, or soil conservation. From time to time
the Programme does undertake occasional in-depth ex post
project and sector evaluation.
The mandate of the United Nations Environment
Programme (UNEP) is to coordinate and catalyse action on
the global environment. Funding for UNEP programmes
comes from the United Nations, trust funds, counterpart
contributions, and voluntary contributions to the
Environment Fund. By far the largest source of funds is the
Environment Fund category. Since contributions to the fund
are voluntary they can vary substantially from year to year.
Contributions decreased in real terms through the early to
mid-1980s, but a recent turn around has seen real
contributions increase by roughly 7% from 1987-89 and by
25% in 1990. With 1990 contributions at just over $50
million the Executive Director has called for the pace to
continue in order that UNEP reach a target of $245 million
by 1995. In 1990 UNEP made commitments of $3.2 million
to support its efforts in biological diversity conservation.
This sum includes work on genetic resources, the
biodiversity convention, biotechnology transfer and
cooperation with NGO conservation initiatives.
The mandate of the UN Food and Agriculture Organization
(FAO) Programme to support development efforts in
fisheries, forests and agriculture means that FAO policies
and activities have a large impact on biodiversity.
Recognising the onslaught of environmental degradation in
developing countries, FAO’s 1989 Governing Conference
added biotechnology and the Tropical Forestry Action Plan
(TFAP) as FAO priority areas. FAO Division directors
meet in working groups on technical environmental matters
including biological diversity. EIA procedures initiated in
1988 are now utilised for evaluating FAO field projects and
projects prepared by the FAO Investment Centre.
Numerous activities undertaken or coordinated by FAO are
designed to mitigate biodiversity loss. FAO is involved in
revising the TFAP, assists UNESCO with the Man and the
Biosphere Programme, and has cooperated with IUCN on
the Caring for the Earth Strategy. Together with UNDP and
512
the World Bank, FAO sponsors the Consultative Group on
International Research (CGIAR) which supports the 13
International Agricultural Research Centers (I[ARCs). While
Green Revolution crop research at these [ARCs is criticised
for leading to loss of on-farm diversity, the International
Board for Plant Genetic Resources (IBPGR) - also an IARC
- is heavily involved in ex situ preservation of genetic
diversity. FAO has also formed a Commission on Plant
Genetic Resources and is involved in developing Regional
Animal Gene Banks.
Biodiversity and environmental initiatives undertakento date
by the major multilaterals indicate that efforts to control
harmful projects and identify beneficial ones, particularly
with regards to biodiversity, are still in their formative
stages. Some organisations have firm guidelines in place;
others are still formalising such procedures. Meanwhile
other agencies prefer a less explicit approach, believing that
concerns over the environment and biodiversity must
become an integral part of the project cycle, instead of an
extra component or evaluation tacked on to the normal
appraisal process.
A key limitation in generating rapid changes surrounds the
capability and availability of staff and consultants. Again
the amount of effort put into retraining varies from one
agency to the next, and from one discipline to the next.
Technical specialists in environmental sciences capable of
carrying out physical EIAs are likely to be more widely
available than environmental economists. As basic and
applied research into the socio-economic impacts of
decrements and increments in environmental quality
improves the tools of the trade, more complete cost-benefit
analysis of projects will be possible.
The Global Environmental Facility
In 1989 at the annual IMF-World Bank Development
Committee meetings, France suggested the creation of a
global fund for encouraging developing countries to
undertake environmental protection activities that provide
benefits to the global community. By November 1990
agreement had been reached by 25 countries that the World
Bank, UNDP and UNEP would cooperate in administering
the Global Environmental Facility (GEF), a mechanism for
distributing concessionary finance for the purpose of
protecting the ‘global commons’.
As of March 31, 1991 twenty-one countries had committed
approximately US$1.4 billion to the fund over a three-year
pilot stage. The Facility is accepting proposals for funding
in four areas:
© protecting the ozone layer
e limiting greenhouse gas emissions
© protecting biodiversity
© protecting international waters.
The Ozone Layer Trust Fund will administer US$160
million earmarked for activities in conjunction with the
Montreal Protocol. Biodiversity and the other two activity
areas will receive funding from the Global Environmental
Trust Fund. The GEF’s mandate with respect to
biodiversity is to preserve specific areas that contribute
goods and services such as harvestable material for
medicines or industrial products, genetic resources for food
production and the regulation of climatic and rainfall
patterns.
Access to GEF funds is limited to countries with GNP of
less than US$4,000 in 1989 and that have UNDP
programmes. In order to differentiate between projects that
meet the GEF mandate and normal development projects
International Aid
Likely candidates for funding are those projects that do not
meet overall rate of return criteria but produce global
environmental benefits and funded projects that with
additional investment could provide such benefits. Projects
should demonstrate that funding cannot be obtained from
other sources such as bilateral and internal sources and they
are not economically viable by normal lending criteria.
Additional qualifications of the projects are the use of
appropriate technology, cost-effectiveness, merit from a
proposals the Bank is developing criteria for GEF projects. global perspective, and consistency with existing
Table 32.6 Global Environment Facility (GEF) biodiversity projects
GEF investment projects
COUNTRY PROJECT TARGET ASSOCIATED FUNDING
PROJECT US$millions
Congo Congo Tropical Forest Lowland rain forest Free-standing 10.00
Preservation
Kenya Lower Tana River Primates Riverine forest IBRD US$30m 6.20
Uganda Gorilla Reserve Bwindi Forest Lowland and montane Free-standing 4.00
forest
Bhutan Trust Fund for Environment Lowland, temperate and Free-standing 10.00
Conservation alpine forests
Laos Wildlife and Protected Areas Lowland and montane IBRD US$10m 5.50
Management forest
Philippines Conservation of Priority Protected 10 high priority protected IBRD US$158m 20.00
Areas areas
Algeria El Kala National Park Wetland IBRD US$30m 12.00
Poland Forest Biodiversity Temperate/montane forest Free-standing 4.00
North Africa New World Screw Worm Wildlife biodiversity FAO Project 9.00
Eradication US$56m - 1991
Brazil National Conservation Units 25 conservation units IBRD US$117m 30.00
Mexico Biodiversity Conservation 20 protected areas IBRD US$30m 30.00
Subtotal 140.70
GEF technical assistance projects
COUNTRY PROJECT DESCRIPTION FUNDING
$millions
East Africa Support for training, research, equipment and institutional development of government, 10.00
university, and NGOs working in protected area management
West/Central Establish a regional TRAFFIC office in Zaire and develop capacity to monitor both legal 1.00
Africa and illegal trade in wildlife
Viet Nam Training/institutional development to prepare a plan for protected areas 3.00
South Pacific Establish and manage 20 conservation areas with threatened biodiversity 8.20
Colombia Assess diversity of the Choco Region through capacity-building research with a view 9.00
to developing plans for protection and sustainable use
Guyana Protect a large tract of rain forest, study the impact of local management 3.00
Amazon Institutional strengthening within the eight members of the Treaty for Amazonian 4.50
Cooperation
Subtotal 38.70
Total GEF biodiversity funding 179.40
513
3. Conservation of Biodiversity
environmental conventions and national environmental
strategies.
Maximum size for freestanding projects is US$10 million.
Project proposals may come from bank or bilateral staff,
government agencies in developing countries and NGOs.
The local UNDP representative must review NGO proposals
with the host government. Projects are forwarded to the
World Bank Regional Environmental Division Chief for
routing to the appropriate desk officer. Bank staff prepare
a three to five page project summary assessing the
compliance of the project with GEF criteria. If the project
looks promising it is then reviewed by a technical panel
which issues a Final Executive Project Summary for
appraisal by the Bank and UNDP and UNEP.
For the fiscal year 1992, the first tranche of funding under
the three year plan, 26 investment and technical assistance
projects are slated for approval at an estimated cost of
US$273 million. Table 32.6 reveals that expected funding
for biodiversity projects totalled US$179 million from the
first tranche. The Bank attributes this to ‘unmet demand’
for financing biodiversity protection. In the remaining years
of the pilot programme the Bank expects to address this
imbalance in the lending portfolio. The share of the initial
GEF devoted to biodiversity projects is expected to reach
US$400 million.
The GEF’s mandate to subsidise the provision of global
benefits by developing countries represents an important
step forward in recognising the distribution of economic
benefits provided by genetic and biological resources.
However, the tone of the projects under consideration by
the GEF is very ‘preservationist.’ As shown in Table 32.6,
despite the variety of approaches employed, almost all the
biodiversity projects are concerned with creating protected
areas or building ‘park’ management capacity. This is a
natural outcome of the GEF’s mandate to fund projects with
low rates of return and large external environmental
benefits. Relatively few of the project briefs include
components that diverge from pure preservation by
encouraging local use and conservation of biodiversity.
However, these activities are considered secondary by the
GEF because of the perception that their benefits are
appropriated locally not internationally. Many consider it
unfortunate that GEF policy contains little overt
acknowledgement that protection and sustainable use of
biodiversity at the local level are often inseparable.
A portion of the GEF portfolio (perhaps through the
proposed small grants window) might be allocated to
research and to projects that encourage the sustainable use
of local diversity with an eye towards external benefits. In
this manner GEF might assist in developing or rekindling
local people’s respect for the benefits of biodiversity and
thereby ensure its continued existence. While a pure
economic analysis of the distribution of benefits from
biodiversity informs the GEF biodiversity strategy, such
analysis overlooks the practical problem that the generation
of external benefits may not be distinct from the generation
of local use benefits.
A final criticism voiced by environmental groups concerns
public access throughout the GEF project cycle. The issue
514
of confidentiality of Bank documents and _ public
participation in project design and evaluation is a common
complaint with regards to Bank projects - not just GEF
biodiversity projects. An interesting conundrum of GEF
financing is that since such projects are ‘environmental’
projects they are exempt from the EIA process under the
1989 EAOD reviewed above. The absence of a detailed
EIA process exacerbates the difficulty of incorporating
public participation, in both developed and developing
countries, into the project selection process.
INTERNATIONAL ASSISTANCE IN FOREST
MANAGEMENT
This section discusses two examples of international
assistance in domestic regulation of resources.
The destruction of tropical forests has for some years been
an issue of considerable concern. It is no longer perceived
as simply a domestic problem for which national remedies
are to be sought, but as a problem of international concern
requiring an international response. A measure of the
importance attached to the problem is the priority given to
it in the 1992 UN Conference on Environment and
Development (UNCED), where it is one of the three
specific topics on the agenda (the others being biodiversity
and climate change).
Widespread concern about global deforestation resulted in
the genesis of two international programmes in 1983. One
of these programmes, the Tropical Forestry Action Plan
(TFAP), was developed by FAO with the assistance of
numerous international organisations and NGOs. The
second programme was the International Tropical Timber
Agreement (ITTA), which was established as a type of
commodities cartel between the governments of tropical
timber producer and consumer countries.
THE TROPICAL FORESTRY ACTION PLAN (TFAP)
TFAP is a programme run by FAO. It is intended to
provide a mechanism whereby international aid efforts could
be better harmonised and coordinated, with a view to
halting the destruction of tropical forests and promoting
their sustainable development. The programme seeks to do
this by helping countries which have tropical forests to
develop national forest management strategies. These
strategies are intended as the basis for increasing investment
in tropical forestry, with the coordinated assistance of aid
programmes from donor countries.
The plan originated from the Committee on Forest
Development in the Tropics (CFDT) - a statutory body of
FAO - which in October 1983 called for the establishment
of ad hoc groups of experts to identify the main problems
in tropical timber production, and the development of action
programmes to address these problems at regional or global
levels. The ultimate result of the various ad hoc meetings
which followed was a five year action programme to
address deforestation issues. The action programme was
divided into five sections: fuel wood and agroforestry; land-
use and upland watersheds; forestry management for
industrial uses; conservation of tropical forest ecosystems;
and strengthening institutions for research training and
education. Each of the five sections contained a list of
recommended actions and investments for a five year action
programme.
The plan was endorsed in June 1985 by CFDT and adopted
by FAO in October 1985 with the formal release of the
TFAP (FAO, 1985). Initially, the means for the
implementation of the recommendationswere not elaborated
upon to any great extent. The plan merely laid out the
principles and recommendations for the guidance of
development assistance agencies, in order to inform them
how aid might be directed to the objective of sustainable
forestry management.
The process required development of an individual national
level TFAP for every country which had tropical forests.
The development of these national TFAPs was a multi-stage
process coordinated by the TFAP unit of FAO. The TFAP
coordination unit is within the Forestry Department of FAO
and is intended to receive technical support from the
organisation as a whole. Funds allocated to the unit from
the FAO regular programme are currently supplemented by
the Multidonor Trust Fund. The process has evolved over
the years and has been implemented in differing ways in
different countries. The development of the procedure has
been carried out by an unofficial body known as the TFAP
Forestry Advisory Group which meets every six months.
The General Terms of Reference for national TFAPs were
outlined in the first meeting of this group in 1985 and these
have been progressively expanded upon at subsequent
meetings (FAO, 1989b).
Despite individual differences in the implementation of the
national TFAPs, their development has involved a series of
basic steps. First, the process is initiated by a request for
assistance from the TFAP coordination unit in the
preparation of a national forest action plan. The next stage
is that FAO or another agency chosen from among the
donors (the World Bank, FINNIDA, CIDA, ODA, etc.) is
identified as the lead agency for the development of that
national TFAP. Then, a review is prepared by the lead
donor agency on the basis of existing information. The
review is sometimes referred to as the ‘issues paper’. This
review is designed to highlight the major problems facing
the forestry sector in the particular country and is used as
a means of identifying sectors of intervention, terms of
reference for consultants, securing participation of NGOs
and local people, and as a basis for the programme and
schedule for the mission. The government of the producer
country concerned then has a chance to review the draft
issues paper. After this, the review is finalised at a meeting
of all parties involved in the forest sector (a meeting
sometimes referred to as ‘Roundtable I’).
The forestry review mission is next set up. This usually
involves foreign consultants, local government officials, and
staff from the lead agency. The review mission then carries
out a forestry sector review over two or three months in the
country for which the national plan is being developed. The
team’s findings are then discussed at a meeting between
Tepresentatives of the government, aid agencies, review
mission team and various concerned NGOs. This meeting
is commonly referred to as ‘Roundtable II’, and the purpose
515
International Aid
of it is to analyse from a technical point of view the various
reports from the forest review mission.
The results of these discussions are then written up as the
national forest action plan. This is then presented at a
national planning seminar held between government officials
and funding agencies to discuss effective implementation of
the plan. This meeting is referred to as ‘Roundtable III’.
Finally, the plan is presented to a wide range of donor
countries and agencies, who can then use it as the basis for
future development assistance to the particular country. The
plan is not legally binding on the donor agencies or
countries and they are free to use only specific parts of the
plan in structuring their future assistance to the country in
question. The objective of the process is to procure
information that will allow donor countries to make more
informed choices concerning the best uses of their aid
monies, in regard to the development of the tropically
forested countries.
To date, no country has completed every step of the entire
process. The latest figures show that a sector review has
been completed in 34 countries and was at different stages
of completion in a further 51 countries. In 24 countries
where the forest sector review had been completed, there
had also been a Roundtable III meeting. In addition, FAO
has received a further 11 requests from national
governments for initiation of the TFAP process, and 10
others are expected to do so in the near future.
The effectiveness of TFAP
The number of countries involved in the TFAP process has
steadily grown since its inception in 1985. At the end of
March 1990, 70 countries which together include 60% of
the world’s remaining tropical forests have become involved
in the TFAP process. However, none have yet completed
the entire process as envisaged by the TFAP guidelines
outlined above. The status of selected national forest plans
is given in Table 32.7 and summarised in Table 32.8.
TFAP has generally been well received by most donor
agencies. More than 40 aid agencies - which together
account for nearly all of the official development assistance
provided to the forestry sector - have collaborated to
support the organisation of over 50 national forest sector
reviews.
Funding commitments to the forestry sector have generally
seen a dramatic increase over the last few years. Total
international developmentassistance increased from US$603
million in 1984 to US$1,095 million in 1988 (FAO, 1989a)
Recently the World Bank has committed itself to tripling
investment in forestry. The ODA has also pledged
£100,000,000 per year to TFAP, and USAID increased
funding of forestry projects from US$50 million in 1988 to
US$72 million in 1989 (Sargent, 1990).
The funds allocated to each of the five TFAP sectors are
indicated in Table 32.9. By contrast, the original TFAP
plan envisaged that the relative funding requirements for
each sector would be: fuel and agroforestry US$1,899
million, land-use on upland watersheds US$1,231 million,
3. Conservation of Biodiversity
Table 32.7 Status of national TFAPs
A. Planning phase completed including Roundtable Ill
LATIN AMERICA/CARIBBEAN
Argentina (National)
Belize (ODA)
Bolivia(UNDP/FAO)
Colombia (Netherlands)
Costa Rica (Netherlands)
Dominican Republic (UNDO/FAO)
Ecuador (National)
Honduras (National)
Jamaica (UNDP/FAO)
Panama (UNDP/FAO)
Peru (CIDA)
Central America (USAID)
B. Forestry sector review completed
LATIN AMERICA/CARIBBEAN
Cuba (National)
Guatemala (USAID)
Guyana (CIDA)
Mexico (National/FAO)
C. Forestry sector review under way
LATIN AMERICA/CARIBBEAN
Chile (FAO/Netherlands)
Haiti (UNDL/FAO)
Nicaragua (SIDA)
Suriname (FAO)
Venezuela (National)
CARICOM (FAO/ODA)
Antigua and Barbuda
Barbados
Dominica
Grenada
Montserrat
St Kitts and Nevis
St Lucia
St Vincent and the Grenadines
Trinidad and Tobago
Amazon Pact (FAO)
AFRICA
Cameroon (UNDP/FAO)
Equatorial Guinea
Ghana (FAO/WB)
Sierra Leone((UNDP/FAO)
Sudan (WB)
Tanzania (FINNIDA)
Zaire (CIDA)
AFRICA
Guinea (France)
Mauritania (UNDP/FAO)
Somalia (UNDP/FAO)
AFRICA
Burkina Faso (GTZ)
Burundi (UNDP/FAO)
Cape Verde (Belgium)
Central African Rep. (WB)
Congo (France)
Cote d’lvoire (FAO/WB)
Ethiopia (WB/UNDP)
Gabon (France)
Gambia
Guinea Bissau (WB/EEC)
Kenya (FINNIDA)
Lesotho (UNDP/FAO)
Madagascar (UNDP/FAO)
Mali (France)
Mozambique (FAO)
Niger (UNDP/FAO)
Nigeria (WB)
Rwanda (ACCT-Canada)
Senegal (UNDP/FAO)
Togo (UNDP/FAO)
Zambia (FINNIDA)
CILSS
SADCC
IGADD
516
ASIA/PACIFIC
Nepal (ADB)
Nepal (ADB)
Papua New Guinea (WB)
Philippines (ADB)
Laos (UNDP/FAO)
Sri Lanka (WB)
Fiji (UNDP/FAO)
ASIA/PACIFIC
Indonesia (WB/FAO)
Malaysia (National)
Viet Nam (UNDP/FAO)
ASIA/PACIFIC
Bangladesh (ADB)
Bhutan (ADB)
India
Pakistan (ADB)
Thailand (FINNIDA/UNDP)
Vanuatu
International Aid
Table 32.7 Status of national TFAPs (continued)
D. TFAP exercise requested
LATIN AMERICA/CARIBBEAN AFRICA ASIA/PACIFIC
El Salvador Angola Myanmar
Paraguay Liberia Solomon Is
Uruguay Mauritius
Uganda
Zimbabwe
E. Preliminary contacts and inquiries
LATIN AMERICA/ CARIBBEAN AFRICA ASIA/PACIFIC
Brazil Chad China
Note: Sub-regional exercises are indicated in italics and are counted separately from countries (except for CARICOM, which is at the same time
a subregional and multi-country exercise) where individual country issues are treated also at national level. The international core support agency
is indicated within brackets. Some countries have completed their planning phase without holding a Roundtable III.
Table 32.8 Summary of national TFAPs
COUNTRIES SUB-REGIONAL
Total of exercises:* 86 6
Africa 37 3
Asia/Pacific Uz
Latin America and Caribbean 32 3
Planning Phase completed 24 -
Sector Review completed 10 tl
Sector Review under way 41 5
Exercise requested 11
Note: * Inquiring countries not included
Table 32.9 Distribution of official development assistance by TFAP fields of action in
1988
FIELDS OF ACTION DONOR DEVELOPMENT UN AGENCIES TOTAL
COUNTRIES BANKS
us$" % us$" % us$" % us$' %
Forestry and Land-use 150.0 27.4 13.9 6.5 50 26.6 213.9 22.6
« Forest-based Industries 92.6 17.0 146.4 68.9 63.8 33.9 302.8 32.0
Fuelwood and Energy 97.9 UP /c) 12.9 6.1 47.2 25.1 158.0 16.7
Conservation 50.3 See 20.0 9.4 ial 7.0 83.5 8.8
Institutions 155.5 28.5 19.4 oe 13.8 7.4 188.7 19.9
Subtotals 631.77 100.0 212.6 100.0 188.0 100.0 1,032.3* 100.0
Source: FAO 1989c. Review of International Cooperation in Tropical Forestry.
Notes: ' In millions. ? Includes undetermined US$85.4 million, 13.5% of total, from Federal Republic of Germany.
industrial forestry US$1,640 million, ecosystem assistance is unclear, as is the extent to which the rate of
conservation US$550 million. TFAP indicated that 20% of deforestation has been slowed down by the programme. The
the funding in each sector should be devoted to overall trend in the problem which the TFAP was
strengthening institutions for research, training and established to combat is more certain: recent FAO statistics
education. These totals represent the sum to be spent over have shown that deforestation for open and closed canopy
the entire five years of the action plan. tropical forests has increased from 11.3 million ha per year
in 1980 to 19.0 million ha per year in 1990 (Collins et al.,
TFAP has been an important instrument for the channelling 1991). No precise data are available for closed forests alone
of aid monies to conservation objectives. The extent to but the trend appears to be similar. As a result of this
which it has actually generated additional developmental continuing rise in the rates of deforestation many NGOs
517
3. Conservation of Biodiversity
have been extremely critical of the TFAP. Several reviews
of the TFAP were carried out in 1990 (Colchester and
Lohmann, 1990; Elliot, 1990; Winterbottom, 1990). All
recommended substantial restructuring of the process. Many
TFAP observers feel that the projects were too frequently
developed from the top down and not from the grass roots
level in areas where deforestation is occurring. This
resulted in a preponderance of foreign experts involved in
each project and not enough input from NGOs. Reviewers
also criticised TFAP for failing to ensure that money spent
in the forestry sector in developing countries is spent
effectively and not with detriment to the environment. It is
also claimed that even though the TFAP was established to
approach the problem of deforestation on a cross-sectoral
basis, it has paid scant attention to non-forest issues. There
have been suggestions that TFAP be given greater
independence either by removing it from the FAO
altogether or by promoting the TFAP from the forest
division and making it a separate division. A process of
review by the architects of the TFAP was instigated at a
meeting arranged by FAO in Geneva last year as a result of
these criticisms. The results of this meeting at the time of
writing are still being negotiated.
Despite the perceived deficiencies of the TFAP, it has acted
as an important mechanism for guiding existing aid toward
more effective investment in regard to the tropical forested
nations.
THE INTERNATIONAL TROPICAL TIMBER
AGREEMENT (ITTA)
At about the same time that the CFDT called upon the FAO
to establish the TFAP, at the UN Conference on Tropical
Timber an agreement on tropical timber was being
negotiated and adopted. The International Tropical Timber
Agreement (ITTA) came into force on 1 April 1985. The
initial term of the agreement was meant to be five years,
but this was extended in 1990 for a further two years. The
ITTA was originally conceived as a commodities agreement
and the initial version, which was drafted by the Japanese,
was based upon the 1979 Rubber Agreement and the 1982
Jute Agreement. However, the final form of the adopted
agreement is unlike previously negotiated commodity
agreements. Typically, commodity agreements are
principally concerned with price control and stabilisation,
and to this end they establish buffer funds and price
manipulation mechanisms. The ITTA, however, developed
into a mechanism much more similar to an agreement for
international development assistance.
This change in emphasis is evident in the preamble to the
agreement. There it states that the parties enter into the
agreement "recognising the importance of and the need for
proper and effective conservation and development of
tropical timber forests with a view to ensuring their
optimum utilisation while maintaining the ecological balance
of the regions concerned and of the biosphere". This
emphasis on sustainable development is also evident in the
objectives of the agreement. The ITTA’s objectives include
the development of the industry, but also the promotion of
research and development with a view to improving forest
management. The purpose of this is to "encourage members
to support and develop industrial tropical timber
518
reforestation and forest management activities" and to
encourage the development of national policies aimed at
sustainable utilisation and conservation of tropical forests
and their genetic resources and at maintaining the ecological
balance in the regions concerned.
The agreement establishes a complex institutional structure
to facilitate the attainment of these objectives. The
administrative structure as a whole is known as the
International Tropical Timber Organization (ITTO) which
is composed of the following elements:
e The International Tropical Timber Council (ITTC)
e Three permanent committees
Committee on Economic Information and Market
Intelligence (PCM)
Committee on Reforestation and Forest Management
(PCF)
Committee on Forest Industry (PCI).
e The Executive Director and staff based in Yokohama,
Japan.
The ITTC acts as the principal political body of the
organisation. The ITTC coordinates the work of the three
permanent committees and carries out all necessary
functions to fulfil the provisions of the ITTA. The Council
is composed of all members of ITTO (i.e. parties to the
ITTA). The ITTC is obliged to hold at least one meeting a
year, although usually two are held - one in Yokohama and
the other in a producer country. The voting scheme used at
these meetings is quite elaborate, and is based upon
achieving balanced representation between industry
producers and consumers (Hypay, 1986).
Permanent committees
Most of the decisions taken by the ITTC are on the basis of
the recommendations of the permanent committees who
function as the operational arm of ITTO (Hypay, 1986).
Participation in each of the permanent committees is open
to all members of the ITTA. As the operational arm of
ITTO these permanent committees play a vital and
important role in the implementation of the objectives of the
ITTA.
The functions of each of the three permanent committees»
are outlined in the ITTA. Apart from normal commodity-
type functions the PCM and the PCF have functions which
would normally be associated with an environmental
protection organisation. For instance, the PCM functions
include "making recommendations to the council on the
need for and nature of appropriate studies on tropical timber
including long term prospects of the international tropical
timber market". The PCF is the key committee for the
implementation of the environmental objectives and its
functions include: reviewing assistance provided at national
and international levels for reforestation and forest
management; encouraging technology transfers for
reforestation and forest management; setting the
requirements and identifying possible sources of financing
for reforestation and forest management. The most
important function of the PCF is "to coordinate and
harmonise these activities for cooperation in the field of
reforestation and forest management with the relevant
activities pursued elsewhere, such as those under FAO,
UNEP, the World Bank, Regional Banks and other
competent organisations". Overall, the permanent
committees are orientated to assisting the general support
role which ITTO plays in this area.
The work of the permanent committee
To date the vast majority of the work of ITTO has been
carried out by these three permanent committees. By the
end of 1990 ITTO had approved 66 projects and
commissioned a further 35 pilot studies for further projects.
The PCF had been delegated the largest number of these
studies and consequently had the largest budget. As of
1 January 1991 this committee was engaged in 19 pre-
project studies and a further 21 current projects. Its
estimated budget for these studies came to over US$12.3
million of which over US$2.5 million had already been
paid. The PCM was engaged in six pre-projects studies and
a further eight current projects. Its estimated budget for
these studies came to over US$2 million of which US$1.3
million had already been paid. The PCI was engaged in
nine pre-projects and a further 19 current projects. Its
estimated budget for these studies came to over US$6.4
million of which over US$2.4 million had been paid.
Table 32.10 outlines the projects which the PCF was then
involved in at the end of 1990. An examination of the
project descriptions show that these closely match the
defined functions of the permanent committee. They also
reflect the idea that this agreement is concerned with more
than the narrow question of maximum price control of a
particular commodity. In fact it appears from the activities
of the permanent committees to date that the question of
price control has not featured very largely on the agenda of
the organisation. Furthermore, from Table 32.10 the
support role envisaged for ITTO in pursuing these
environmental objectives is evident in both the nature of the
projects and the fact that in only five of the 21 current
projects is ITTO the implementing or lead agency.
Apart from sponsoring these projects, the other major
initiative of ITTO has been a programme to assist countries
in the development of management procedures to direct
timber production in tropical forests toward sustainability.
The year 2000 was established as the target date for the
achievement of sustainable management of tropical forests
worldwide. This object has been pursued through the
establishment of forestry standards for the sustainable
Management of natural tropical forests for timber
production. These were drawn up by the PCF and adopted
by the ITTC at the 7th session of the Council in May 1990.
These standards contain a set of 41 principles and 36
possible actions. They cover considerations ranging from
general policy to particular aspects of forestry operations.
The general principles involved include the establishment of
national forestry inventories and a permanent forest estate.
They also recommend examination of forest lands
ownership, and the establishment of separate institutions for
the management of the forest estate in each country. In
addition to these guidelines ITTO is presently developing
another set of guidelines on biodiversity, known as ‘The
ITTO Guidelines on the Conservation of Biological
Diversity in Tropical Production Forests’. The objective of
519
International Aid
these guidelines is to "optimise the contribution of these
forests to the conservation of biological diversity that is
consistent with ... the sustainable production of timber and
other products". At the time of writing these guidelines,
although accepted by the PCF committee, have not gained
political support at the ITTC level and are unlikely to be
acted upon by the ITTC before UNCED (Anon., 1991).
In addition to the development of these standards, ITTO is
also generating financial assistance for producer countries,
both directly from its own funds and indirectly through the
solicitation of contributions from consumer countries. This
assistance is usually employed to help countries in their
incorporation of these guidelines into national policy and
legislation.
Conclusion
From the above overview of TFAP and ITTA, it is evident
there are a number of similarities between the two
organisations. This is to be expected as they are both
international responses to the problem of deforestation
represents a significant global problem. One programme
originated out of the international resource community, and
the other out of the industry itself.
The stated purpose of TFAP is to harmonise and coordinate
actions in the tropical forest sector so that tropical forests
can be used by mankind on a sustainable basis. Similarly,
the stated purpose of ITTO is "to develop proper and
effective conservation and development of Tropical Forests
with a view to ensuring their optimum utilisation while
maintaining the ecological balance of the regions concerned
and of the biosphere." ITTO states that it wants to move
worldwide production to a sustainable basis by the year
2000.
Despite the fact that the objectives of the two organisations
are similar, the way in which both organisations set out to
achieve these objectives is ostensibly quite different; this is
because of the origins of the two programmes and the
difference in their legal structure.
The TFAP is technically a policy which lacks any legal
content. From a legal point of view it is no more than an
information document which organisations working in the
area may choose to adopt to guide their decisions. The
funds dispensed under the programme do not pass through
the TFAP; instead, they flow directly from the donor to the
recipient. At the discussions about the implementation of a
national TFAP (Roundtable II), if a donor can be found for
a particular aspect of the plan or the entire National TFAP,
then that donor will make arrangements for the provision of
funds to the recipient. Another consequence of the legal
structure of the TFAP is that few of the activities involved
in the development of a national TFAP are actually carried
out in the name of the TFAP; they are instead coordinated
and conducted in the name of the lead agency. In sum, the
TFAP is only a name associated with a large number of
separate agreements and relationships; it has no structure of
its own.
ITTO, on the other hand, is a legally constituted
organisation which does have its own legal personality,
Conservation of Biodiversity
3:
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52
3. Conservation of Biodiversity
headquarters, staff and budget. ITTO carries out many
activities in its own name and raises funds and uses its
resources in its own name.
An important reason for the different approaches is the
different origins of the two programmes: the TFAP was
developed by FAO and various NGOs active in the tropical
forest sector whereas ITTO is a product of the UN
Conference on Trade and Development (UNCTAD). Whilst
TFAP has approached the problem from the coordination of
development assistance programmes, ITTO has approached
it much more from a trade point of view. Thus TFAP is
meant to approach the problem in a cross-sectoral way and
has sought to include as many parties as possible in the
process. It has traditionally attempted to do this through the
Roundtables, by increasing the importance of the forestry
sector and by coordinating the various development
assistance programmes that have some impact on the forest
sector in the relevant country. On the other hand ITTO is
not cross-sectoral and is concerned simply with the forest
estate of the producing countries. This different approach is
seen in the nature of the projects ITTO is involved in,
which on the whole tend to be concerned with quite specific
problems in tropical timber production.
Yet, despite these ostensible differences in approach, the
work of the two programmes is very similar. The defined
functions of the PCF are very similar to the fields of action
of the TFAP. For instance the TFAP calls for "the
protection and management of natural forests" along with
"accelerated industrial reafforestation" and ITTO has
established a permanent committee whose main purpose is
to "promote better forest management and reafforestation’ -
the PCF. TFAP states that "financial incentives are needed
to encourage investment in reafforestation and forest
management" and the ITTA empowers the PCF to "identify
all possible sources of financing for reforestation and forest
management". Both programmes make extensive reference
to promoting training, research and education in the area.
Both encourage greater transfer of technology. The
guidelines developed by ITTO closely resemble many
national TFAPs.
This overlap between the two organisations results in some
duplication of effort and illustrates a lack of direct
cooperation between the TFAP and ITTO. This is perhaps
one of the more easily correctable failings of international
efforts to assist in the regulation of this domestic resource.
DEBT PURCHASE
The debt purchase discussed here covers a specific form of
debt-equity conversion, widely termed a ‘debt-for-nature’
swap. Other types of debt-equity conversions such as ‘debt-
for-development’ and ‘debt-for-child’ also occur. The
essential aim of all these types of instruments is to convert
the external debt of a developing country into a domestic
obligation to support a specific programme. Table 32.11
details the debt-for-nature agreements which have been
established so far. Although no two debt-for-nature swaps
so far negotiated have been identical, the basic structure
used in each case is similar.
The first step is that an international conservation group
must raise funds in order to ‘purchase’ a debtor country’s
tw
foreign debt. Private banks are usually reluctant to make
outright donations of the debt they hold, even though in
some countries like the USA such a donation is given a
favourable tax treatment. Funds are usually secured from
either the international conservation group’s own resources
or from donations from private individuals or bilateral aid
agencies.
The funds raised are used to purchase the country’s external
debt on the secondary market at a fraction of the theoretical
or face value of the debt. The ‘secondary market’ is a term
used to describe the process whereby the original creditor
of the debtor country sells on part or all of the debt to
another institution. This trading can happen many times and
is so prevalent that rarely will the bank who arranged the
original loan retain anything but a small portion of the
original debt. Table 32.11 shows that typically the country’s
debt has been purchased at between 15-30% of its face
value. This discounting is because of a low expectation of
total repayment by the debtor countries; the amount of the
discount is proportional to the expectation of repayment.
Once the external debt is acquired, the environmental
organisation will enter into negotiations with the debtor
country to fix a favourable rate for the conversion of the
external debt from the foreign currency in which the debt
is denominated to the local currency of the debtor country.
This rate will usually be somewhere between the local
currency value of the debt and the local currency value of
the price the environmental organisation paid for the debt
on the secondary market. The price that is negotiated is
referred to as the redemption price. Most commonly the
redemption price is 100% of the face value of the acquired
debt but in some instances it may be no more than the
discount value of the acquired debt.
Lastly, the debtor country’s government issues a financial
instrument, typically a government bond, denominated in
local currency in an amount equal to the redemption price
of the debt. These bonds are then used to finance projects
in the debtor country through local organisations.
A debt-for-nature agreement is often described as one in
which all the parties involved stand to gain something.
The international conservation group is able to increase the
spending power of its usually limited financial resources
because of leverage provided by the difference between the
redemption price and the discount rate of the purchased
external debt. The group is also able by this method to
influence conservation policy in countries where normally
they have little impact. For the international conservation
group there are also numerous collateral benefits, such as
the relationship created between the parties involved (e.g.
the local Ministry of Environment, Ministry of Finance,
local conservation groups and creditor banks).
For the investors and the institutions who hold the country’s
debt there are direct benefits associated with having the
extra purchases in the secondary markets, such as
increasing both liquidity of the market and the price of the
discounted debt.
For the debtor countries there are considerable financial
benefits to be gained from the debt-for-nature agreement.
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3. Conservation of Biodiversity
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525
3. Conservation of Biodiversity
Firstly, it reduces the debt owed by the country by an
amount equal to the difference between the redemption price
and the face value (although this can be illusory because, as
noted above, countries can themselves step into the
secondary market and purchase the discounted debt;
therefore the redemption price becomes equal to the
discount price. Most importantly, the country reduces its
foreign debt by this mechanism. This in turn reduces the
often crippling need to raise foreign currency to service the
country’s existing debt and thereby helps its external
balance of payments. Another important political benefit is
that the government of the debtor country will be able to
control the donation made by the international conservation
group, whereas if the donation had taken place directly
from the international conservation group to the local
conservation group the government would have less control.
Notwithstanding the potential benefits of debt-for-nature
agreements, there are several problems which limit their
usefulness.
Several countries, the Brazilian government foremost
amongst them, have stated that the environmental conditions
which are associated with such agreements are an
imposition on the foreign sovereignty of the debtor nations
(and contrary to the UN General Assembly Resolution 1803
on Natural Resources). This curtailment of sovereignty is
supposedly manifest in two ways. Firstly, vesting in foreign
creditors control over the debtor country’s land and natural
resources is in a sense equivalent to selling these resources
to the outside interests. Secondly, it is claimed that the
debt-for-nature agreements facilitate the imposition of
foreign projects and values, and influence local policy in
conservation projects in a manner which is more beneficial
to outside interests than to local interests.
These fears seem largely unsupported by the facts of each
debt-for-nature agreement established so far. None of the
debt-for-nature agreements has entailed the transfer of
ownership or control to foreign creditors of any sort
including the international conservation group. Rather, in
each instance the debt-for-nature arrangements have
transferred control of the debt from foreign interests to
local concerns. The involvement of the international
conservation group does unquestionably to some extent
impose foreign values; however, because of the need for
complete cooperation of the debtor government, outside
interests are subject to veto by local concerns. Most debt-
for-nature agreements have been proposed by local groups
within the debtor nations as a means of increasing the
effectiveness of their own projects. Consequently any
foreign influence exerted by the international conservation
group is more in the nature of a positive exchange of ideas
rather than direct imposition of inappropriate values which
critics suspect.
Furthermore, debt-for-nature agreements are too small to
have the consequences feared by critics. As can be seen
from Table 32.11, the total amount of foreign debt which
has been retired by means of debt-for-nature agreements is
less than US$100 million, compared to the total debt owed
by the developing world of an estimated $1.3 trillion. The
insignificant size of these debt-for-nature agreements,
however, does raise another point of genuine concern which
526
is thatany attention given to debt-for-nature agreements will
direct attention away from solutions on a more meaningful
scale to the problem of debt in less developed countries.
Finally, it is argued that the debt-for-nature mechanism, if
it were to be implemented on any significant scale, would
have an inflationary effect on the debtor country’s
economy. However, this will only be the case where the
financial instruments used by the government to pay the
redemption price negotiated by the international
conservation group is local currency.
It should also be noted that because the attractiveness of
debt-for-nature swaps for the international conservation
group is the fact that the debtor countries’ debt trades at a
deep discount in the secondary market, this instrument
cannot be used to make significant inroads into the foreign
debt of a debtor country. For as soon as any significant
amount of debt for a particular country starts to be
purchased by parties wishing to use debt-equity instruments
to fund projects in the country, this will drive the
discounted price of the debt up, and it therefore becomes of
diminished attraction. Indeed, as recent months have shown,
the uninspiring economic situation in much of Latin
America, for example, has seen a dramatic rise in the price
at which debts are being traded on secondary markets.
These criticisms, however, do nothing to detract from the
main importance of the mechanism. Debt-for-nature swap
agreements should not be seen as a way of reducing the
foreign debt of a country but rather as a means to help
develop the promotion of environmental ideas and projects
within a country through the local environmental
programmes and groups in that country.
References
Abramovitz, J. 1991. Investing in Biological Diversity: U.S. research
and conservation efforts in developing countries. World Resources
Institute, Washington.
Anon. 1991. Report of the Working Group on Guidelines for the
Conservation of Biological Diversity in Tropical Production
Forests. International Tropical Timber Council. Eleventh Session
28 November-4 December 1991. Yokohama.
Colchester, M. and Lohmann, L. 1990. The Tropical Forestry Action
Plan: what progress? World Rainforest Movement and The
Ecologist. Penang, and Sturminster Newton, Dorset.
Collins, N.M., Sayer, J.A. and Whitmore, T.C. 1991. The
Conservation Atlas of Tropical Forests: Asia and the Pacific.
Macmillan Press, London, UK, in collaboration with IUCN, Gland,
Switzerland.
DAC 1990. Development Co-operation: efforts and policies of the
members of the Development Assistance Committee. OECD, Paris.
Dogsé, P. and Droste, B. 1990. Debt-for-nature exchanges and
biosphere reserves: experiences and potential. MAB Digest 6.
Unesco, Paris.
Elliott, C. 1990. The Tropical Forestry Action Plan. World Wide Fund
for Nature International, Gland, Switzerland.
FAO 1985. Tropical Forestry Action Plan. Committee for Forest
Development in the Tropics. Rome.
FAO 1989a. Committee on Forest Development in the Tropics. Ninth
Session. Papers supporting the Agenda.
FAO 1989b. Guidelines for Implementation of the Tropical Forestry
Action Plan at the Country Level. Forestry Department. Rome.
FAO 1989c. Review of International Cooperation in Tropical Forestry.
Hypay, T. 1986. The International Tropical Timber Agreement. Its
prospects for tropical timber trade, development and forest
management. YUCN/IIED, London. 18pp.
Sargent, C. 1990. Defining the Issues: some thoughts and
recommendations on recent critical comments of the TFAP. ITED.
13pp.
Silva, L. de 1982. Development Aid: a guide to facts and issues. Third
World Forum and UN NGLS, Geneva.
UNEP 1990. UNEP Profile. UNEP, Nairobi.
Winterbottom, R. 1990. Taking Stock: the Tropical Forestry Action
Plan after five years. World Resources Institute, Washington, DC.
527
International Aid
World Bank 1989. Operational
Environmental Assessment.
Directive 4.00, Annex A:
Authors as follows: International development assistance,
Bruce Aylward (LEEC/IIED); International assistance in
forest management, Sam Johnston; Debt purchase, Victoria
Drake.
3. Conservation and Management of Biodiversity
33. MANAGEMENT OF INTERNATIONAL RESOURCES
National boundaries do not enclose all the world’s
biological diversity; the high seas, the deep sea bed and
Antarctica all contain natural resources, some of great
interest or economic importance. Management of
biodiversity in such areas can, by definition, only be
achieved by means of international measures.
This chapter will discuss measures taken by the
international community for biodiversity conservation in
international areas, and highlight some of their strengths
and weaknesses.
Use of natural resources in such areas is characterised by
over-exploitation and resource depletion; this is typified by
the whaling and sealing industries and several fisheries.
These are classic examples of over-exploitation as a result
of unrestricted access by all users, none of which has any
incentive to limit extraction in the interests of long-term
sustainability. With open-access resources the benefits of
forbearance do not accrue to those who exercise it but
rather to other users, who simply end up with a greater
percentage of the market. Economically-effective use of
capital also demands that exploitation occurs sooner rather
than later.
A common response to these problems has been the
establishment of an international commission mandated to
control the use of a particular resource. The International
Whaling Commission is one example, and we discuss below
some other commissions concerned with international
fisheries. These commissions typically use two different
types of measure in an attempt to conserve their resource:
setting of quotas and the setting of minimum standards
which operators must adhere to.
Most of these international commissions have had only
limited success in controlling over-exploitation. Some of the
more common reasons for the failure of these commissions
are: lack of finances, lack of political consensus, lack of
scientific information about the resource and lack of power
to monitor compliance with the controls established.
On the other hand, the development of the Antarctic Treaty
System has been a comparatively successful venture in
international resource management. The process of
developing the Antarctic legal system has occurred over a
period of decades, but it has been a consistent progression
from a very broad and uncertain regulatory system to one
that is now reasonably well-defined.
There is some tendency for change in the way ownership of
international resources is perceived; they can be regarded
as belonging not to those who appropriate them, but to
mankind as a whole. One implication of this change is that
international resources should be managed for all of
mankind and not simply for those who have the ability to
appropriate them. While this much is not especially
contentious, many international statements have been made
to the effect that biodiversity in general is the common
heritage of mankind. The policy and legal implications that
this view has on resources within national boundaries are
complex and contentious, and have provided a major subject
528
of debate in discussions preparatory to the proposed
Biodiversity Convention.
INTERNATIONAL FISHERIES MANAGEMENT
COMMISSIONS
The need for regulation
Fisheries have traditionally been regarded as common
property, and so open to all without restriction. The typical
pattern of use of common property resources involves
increasing production beyond the point of sustainability until
production declines (Hardin, 1969). This is because the
economic imperatives of common property are to harvest
the resource before someone else does, forgoing investment
that would improve productivity of the resource.
This pattern of common property exploitation has occurred
repeatedly in fisheries. The decline of stocks of herring and
mackerel in the north-east Atlantic, the King Crab in the
north-east Pacific, Yellow-fin Tuna in the eastern Pacific
and Blue-fin Tuna in the south-west Pacific oceans are cases
in point (Brown and Crutchfield, 1981). The human
consequences of stock decline include smaller catch per unit
effort, excess capacity of fishing equipment (large powerful
vessels with sophisticated equipment) and reduced income
for fishermen.
It is widely recognised that techniques must be introduced
to manage stocks more efficiently, minimise costs and
improve distribution of fishery benefits. Fisheries
commissions have been established to perform these tasks.
Legal parameters for fisheries commissions
Overfishing became a recognised international problem
initially for a few stocks in the North Sea in the late 19th
century. However, international regulation is essentially a
20th century phenomenon (Underdal, 1980) and was not
widespread before the first half of the 20th century.
Legal arrangements for cooperative managementof fisheries
can be regarded as falling into two periods, with the
division between them being the rising importance of the
Exclusive Economic Zone (EEZ) in the mid-1970s. This
brought most traditional high seas fisheries under the
jurisdiction of the coastal State, because most fish stocks
are found within the 200 nautical mile (nm) EEZ. Until that
time, coastal States had jurisdiction over only their
territorial seas, extending 3-12nm from the coast. Beyond
that, all States enjoyed open access to fisheries in the high
seas. The proclamation of EEZs by coastal States extends
their jurisdiction to more than half the total area of the
world’s seas, and the great majority of the customarily
exploited living marine resources. Much variation in the
catch patterns of particular countries has, therefore, been
because of the new authority of coastal States over adjacent
fisheries, which has enabled them to exclude or control
long-range water fleets (Brown and Crutchfield, 1981).
Prior to the declaration of EEZs, there was an urgent need
for cooperative management of high seas fish stocks. A
range of treaties regulating the harvest of fisheries made
their appearance early this century. The USA and Canada
agreed on conservation treaties for halibut and salmon in the
north-east Pacific, and European regional agreements were
concluded for the Baltic and North seas and the northern
Atlantic.
By the late 1950s fishing methods were changing rapidly.
The use of sonar, mother ships with on-board processing
facilities and purse seining were prominent (Knight, 1975).
Fishing patterns were also altered as distant water fleets of
developed countries, in particular those of Japan and the
former USSR, expanded to all oceans and placed heavy
pressures on fish stocks.
The 1958 Convention on Fishing and Conservation of
Living Marine Resources became the first global fisheries
management agreement. It placed an obligation upon its
participating States to cooperate in adopting conservation
measures. The intention of ‘conservation of living
resources’ was solely to render possible the ‘maximum
sustainable yield’ (MSY) from those resources so as to
secure the maximum supply of food and goods for human
consumption. The MSY approach has been criticised for
failing to deal with fluctuations in stock size caused by a
multitude of variables other than catch size, and because it
ignores the fates of associated and dependent species.
Following the global 1958 Convention, several new regional
conventions were concluded for the Atlantic, Baltic and
Black seas. The next global approach to fisheries
conservation to be adopted was the 1982 UN Convention on
the Law of the Sea (UNCLOS). This convention included
the obligation on participating states to conserve fishery
resources in areas under their jurisdiction and to cooperate
in their conservation beyond national jurisdiction. It also
incorporated formal international recognition of the EEZ
concept.
Many fish stocks are highly migratory and relatively few
important commercial species remain within only one EEZ
(Brown and Crutchfield, 1981). Therefore, the creation of
EEZs has not eliminated the need for international
institutions to facilitate cooperation in the management of
fisheries but has simply redefined their role and reduced
their independence. The need for international commissions
is recognised in Article 63 of UNCLOS, which requires
States to form regional or sub-regional organisations in
order to foster cooperation between them in the
conservation of the marine living resources.
UNCLOS forms a very loose and inadequate framework for
decision-making by regional fisheries organisations. It
requires that States maintain harvested species at population
levels sufficient to produce an MSY, as qualified by
relevant environmental and economic factors, and taking
into account the effects on associated and dependent species
with a view to preventing them from being threatened with
extinction. States are also to ensure that conservation
measures do not discriminate against the fishermen of any
State (a constraint which may be difficult to meet in
practice).
The MSY approach may be contrasted with the much wider
parameters adopted in the 1980 Convention on the
$29
Management of International Resources
Conservation of Antarctic Living Marine Resources
(CCALMR). It requires the "prevention of decrease in the
size of harvested population to levels below those which
ensure its stable recruitment", "maintenance of the
ecological relationships between harvested, dependent and
related populations", and "prevention of changes or
minimization of risks of changes in the marine ecosystem
which are not potentially reversible in two or three decades"
(Article II). This approach is better suited to integrated
management and conservation of ecosystems.
Decisions on the international management of fisheries are
primarily concerned with the maintenance of the resource
base but also need to reconcile many conflicting national
and economic interests. These include maintenance of the
industry’s profitability and of the welfare of fishing
communities, and equitable distribution of wealth between
competing fishing States. The political sensitivities inherent
in these interests have often resulted in international
fisheries bodies being limited to very narrow and
uncontentious mandates, such as the gathering and
dissemination of information on fish stocks and fisheries
technologies. Examples include informational bodies
established by the UN Food and Agriculture Organization,
such as the Indo-Pacific Fisheries Commission, the Western
Central Atlantic Fisheries Commission and the General
Fisheries Council for the Mediterranean.
Smaller bodies established by the participating nations
themselves tend more often to have management powers.
The following case studies of international fisheries
commissions examine the effectiveness of a few such
bodies.
European Common Fisheries
Jurisdiction
The European Community (EC) Common Fisheries Policy
(CFP) evolved in the 1970s as a means of ensuring equal
access by EC Member States to each other’s fishing
grounds. It applies in the EEZs along the North Sea and
Atlantic coastlines of Member States and in certain areas of
the west Atlantic, Skagerrak, Kattegat and Baltic Sea
(Farnell and Elles, 1984).
Administration
The CFP is implemented by the EC. In particular, the
Fisheries Council (constituted by Fisheries Ministers of the
Member States) acts as the legislative organ and the
Commission Directorate-General for Fisheriesygcts as the
executive organ and enacts delegated legislation. A range of
minor committees, mostly concerned with the provision of
information, service these bodies. The Fisheries Council
makes decisions on the long-term availability and
distribution of fishery resources and on annual management
matters, such as fishing restrictions, monitoring and
enforcement. It votes in accordance with the usual EC
procedures.
Member States can impose their own conservation measures
only in relation to strictly local fisheries affecting their own
citizens. They can impose unilateral conservation measures
affecting fishermen from other EC States only where fishing
grounds within their jurisdiction are seriously threatened
3. Conservation and Management of Biodiversity
and any damage would be difficult to repair. A range of
other constraints are imposed upon such unilateral measures
and they must be submitted to the Commission to confirm,
amend or cancel. It is apparent that a great deal of
exclusive management power rests with the EC.
Information
The provision of reliable information is crucial to the
operation of any fishery commission. However, they are
extremely difficult and expensive to obtain because of
inadequate scientific knowledge of fishery dynamics. The
EC therefore relies largely on a system of voluntary
reporting. Fishermen must keep logbooks, and make fish
landing and trans-shipment declarations. However, the
voluntary reporting system is flawed as fishermen do not
see it as being in their interests to make declarations which
will ultimately lead to restrictions being placed upon their
activities. The information provided is therefore often
inaccurate. For example, fishermen landed at least 50%
more cod and sole than they were permitted to in 1989.
Independent research is at present conducted for the EC by
the Advisory Committee on Fisheries Management of the
International Council for Exploration of the Seas.
Fleet capacity containment
If the capacity of a fishing fleet does not exceed the
sustainable yield of fish, no restrictions on the fishing effort
of the fleet would be required in order to conserve fish.
Over-capacity is, then, a root problem for fisheries
management. The EC fleet capacity is currently 40% in
excess of available fishing opportunities (Rose, 1991).
This problem is being dealt with by a 10-year scheme of
“Multi-Annual Guidance Programmes’ (MAGPs) to
restructure the EC fleet. Each Member State must present
to the Commission for its approval two five-year
programmes to adjust its fleet capacity, and then report
back annually on implementation. Aid, in the form of a
70% reimbursement, is provided to Member States for the
temporary or permanent withdrawal of vessels from service.
The MAGPs also include provisions for building and
modernising vessels, seeking development of aquaculture in
the EC, exploratory fishing and joint ventures outside the
EC and the development of markets for surplus or
underfished species.
The MAGPs are not very effective as, in countries where
capacity is greatly in excess of fishing opportunities,
significant capacity reduction is not being achieved. As a
consequence, in October 1990 the EC Commission froze
MAGPs3’ grants for construction of new vessels in the UK,
Ireland, the Netherlands and Greece. The MAGPs would be
more effective if they were more tightly regulated.
The EC has recently created a central register for fishing
vessels which includes details of vessel capacity. The use of
a central register creates opportunities for a coordinated
system of EC vessel licences entailing capacity quotas and
penalty systems. Such capacity quotas are already in place
in relation to Spanish and Portuguese fishermen and waters
but remain politically unacceptable across the whole EC
fleet. Incentives for selective fishing methods and
disincentives for non-selective methods could also be
introduced into the MAGPs and, ultimately, an EC
licensing system (Rose, 1991).
530
Fishing restrictions
Restrictions on fishing effort include restrictions on access
to specified areas or during certain seasons, limits on
catches of particular species and undersized fish, and
restrictions on the use of certain gear and on the allowable
end use of some fish. No single restriction is sufficient to
manage a fishery adequately, and usually a mix of measures
is adopted, as is the case in the EC.
Total allowable catch
This is fixed by the Council each year, and quotas are
distributed between the Member States. This is the
foundation of EC fisheries conservation measures. Catch
landings are monitored and reported by Member States and
fishing activities are halted when the quota has been used
up. States are able to trade quotas and obliged to
compensate for the illegal use of another’s quota. However,
this is an essentially quantitative and bureaucratic method of
conservation and is very inefficient. It encourages fishermen
to get ahead of competitors in using up the quota and
requires close monitoring and enforcement in order to
prevent false understatement of landings. Fishermen are
legally obliged to discard fish caught in excess of the quotas
(Rose, 1991).
Minimum size restrictions are imposed on certain protected
species. However, fish below the minimum size must be
thrown back into the water, resulting in unnecessary
wastage. The existing restrictions follow rather than prevent
the catch of protected species and close surveillance is
needed to enforce them.
More effective regulatory measures control fishing
techniques rather than catches. For example, the EC applies
area access restrictions within 12nm of the coast, and a
licensing system for access to areas where there are species
of special importance. These measures are sometimes
supplemented by season and duration restrictions. For
example, the EC has required vessels engaged in certain
fisheries to lay up in port for 10 consecutive days each
month (Rose, 1991). These approaches have negative
economic impact, as boats, equipment and labour become
under-used (Keen, 1988).
Other management measures include gear restrictions and
special licensing for some fishing activities. Gear
restrictions (such as prohibiting the use of guns or
explosives) may be more effective in ensuring that certain
fish are not landed in the first place (Rose, 1991).
Enforcement
The mix of fishing restrictions adopted by the EC requires
a high degree of cooperation from fishermen or else close
surveillance and firm enforcement. These are generally
lacking. The Commission does not have independent
monitoring powers but relies on Member States. These are
obliged to report annually on their inspections at sea and in
port and on the warnings, prosecutions and penalties which
result. Yet national authorities often fail to ensure that
conservation measures are implemented, partly because of
the inadequacy of staff and facilities and also the difficulty
of obtaining evidence.
Unfortunately, the CFP is failing to meet its conservation
goals: 75% of fish stock within the area is exploited at
unsustainably high levels (Rose, 1991). The Commissioner
for Fisheries has even threatened to abandon EC control
over fisheries management if Member States continue to
block the Commission’s management efforts. Although it is
clear that the situation would be worse if the stocks were
unregulated, doubts must be raised as to whether they
would be better managed unilaterally under national
jurisdiction.
North East Atlantic Fisheries Commission
Administration
The North East Atlantic Fisheries Commission (NEAFC)
came into being in 1963, and applies to parts of the Arctic
and Atlantic Oceans. It provided the framework for most
international fisheries regulations in the area until 1977.
By mid-1976, it had 16 Member States, each having two
commissioners, and its principal task was to recommend
conservation measures to ensure rational exploitation of
various stocks. Recommendations were only binding on
States which did not object to them within a certain period.
Application and enforcement were left to each Member
State (Underdal, 1980). Clearly the participation of all
Significant groups was necessary for a measure to be
effective; accordingly, the objection procedure had the
effect of encouraging the development of the least ambitious
management programme.
Catch limits
To establish a total allowable catch (TAC) required the
consent of all Member States, and the approval of two-
thirds of the delegations. This was not obtained until 1974
and by then some stocks were severely depleted. Until this
time, the NEAFC simply made extensions and modifications
to the mesh size and minimum landing size provisions. The
TACs were not effective when set, as they were not
adequately enforced and had to be reset each year by
bargaining between the Members (Mason, 1979).
This usually resulted in the setting of such generous TACs
that sacrifices were not necessary to contain the catches
within them. Table 33.1 shows that in 1975 the total catch
for 11 out of 15 stocks did not reach 90% of the TAC and
in 1976 catches from 8 out of 15 stocks did not reach 90%
of the TAC. The total catch in 1975 was only 84% of the
NEAFC TAC. In many cases the TAC adopted by the
NEAFC exceeded the highest TAC proposed by any party.
This was because the Parties’ inability to resolve arguments
over individual allocations was often eventually resolved by
simply raising the TAC. Table 33.2 indicates this process
for 1975.
Quotas
In theory, the power to impose quotas gave the NEAFC
every possible option to redistribute the TAC between
Members. However, the quota did not cause redistributions
Significantly larger than normal fluctuations under an
unregulated market. Quotas tended to be influenced by
arguments based on rights (i.e. territorial or historical use
of the resource) or on a concept of what is fair, taking into
account needs and responsibilities. As overall economic
power tended not to be a basis for argument, redistribution
of quota tended to move from those with a large catch to
531
Management of International Resources
Table 33.1 National catches as per
cent of NEAFC TACs
STOCK TOTAL CATCHES AS %
OF NEAFC TACs
1975 1976
Arcto-Norwegian Cod 98 101
North Sea Cod 80 89
North Sea Haddock 67 100
North Sea Whiting 81 101
North Sea Plaice 86 106
North Sea Sole 146 111
North Sea Sprat = 95
North Sea Herring 69 is
Herring w.o. Scotland 91 78
Celtic Sea Herring 55 54
Irish Sea Plaice 80 79
Irish Sea Sole 85 83
English Channel Plaice 87 77
English Channel Sole 95 116
Bristol Channel Plaice 59 47
Bristol Channel Sole 81 74
AVERAGE 84 95
Source: NEAFC reports.
Notes: In the catch statistics used as the source for this table the
former USSR catches of coastal cod are included in the figures for
Arcto-Norwegian Cod. For this reason the USSR quota of coastal cod
has been added to its regular quota and to the TAC in the calculation
of this table. The first quota regulations for the North Sea Herring
fishery applied to the period July 1974 through June 1975. Available
catch statistics follow calendar year.
those with smaller claims. Once a quota was agreed on, this
provided a strong precedent for future decision as there was
no precise or generally accepted formula for distributing
TACs, which were not mentioned in the treaty. Cutbacks in
quotas were usually proportional, so as to avoid
redistribution. A Member usually advocated those
arguments which produced the most favourable distribution
for itself. Consequently, 86% of proponents’ schemes gave
proponents a higher proportion of the TAC than did any
other proposal (Underdal, 1980).
The operation of the NEAFC can be most simply examined
through case studies. The most difficult issue it faced was
the management of the North Sea Herring.
In the late 1950s to early 1960s, scientists were concerned
at the decline of the herring. The catch dropped from
225,000 tonnes in 1955 to 45,000 tonnes in 1963, but the
scientific evidence of overfishing was not conclusive. The
NEAFC was slow to respond, largely because of the self-
interest shown by States such as Denmark (Underdal,
1980). It could only get agreement on more research and a
study group was formed in 1969.
In 1970 conservative regulations were passed. Agreement
was made possible by the continued decline of catches and
stock, as it was by then clear that overfishing was a
principal cause. The regulations were ‘formally neutral’, in
that they did not directly regulate individual State quotas.
They concerned limitations on mesh size and landing size,
3. Conservation and Management of Biodiversity
Table 33.2 TAC decisions by NEAFC in relation to initial proposals
STOCK FIRST QUOTA REGULATION
Lowest Highest
Proposal Proposal
Arcto-Norwegian Cod 510 1165
North Sea Cod 220 230
North Sea Haddock 220 260
North Sea Whiting 120 190
North Sea Plaice 115 125/130
North Sea Sole 6 10
North Sea Herring 310 424
Herring w.o. Scotland 156 170/200
Celtic Sea Herring 25 30
North Sea Sprat 300 No TAC
SECOND QUOTA REGULATION
Decision Lowest Highest Decision
Proposal Proposal
810 700/800 800/900 810
236 210 236 1236
275 150 ?155 1206.25
189 160 189 189
126 85 ; 100/105 99.9
12.5 8 12.5 12.5
494 200 ? 2254
205 - < a.
32 719 23 25
589 - = =
Source: NEAFC reports and summary records, December 1973-November 1975. All figures are in 000s tonnes.
Notes: ? figure uncertain. - information not available. 1 The USSR delegation indicated indifference between the lowest and the highest TAC
suggested by the LC (SR, Nov 1975: 1/8). ? The discussion at the 13th annual meeting started with the idea of a 12 month allocation, but soon
developed into a discussion on a regulation covering 18 months. For this reason some of the TAC proposals are hardly comparable. 3 In the
discussion about the second quota regulation, regulations for both 1975 and 1976 were considered, the TAC for one year depending on the TAC
for the other.
and season closures in 1971-1974, which had little effect,
because of the exemptions from them. Depletion continued
and the NEAFC requested power to limit the amount of
catch and effort and to allocate quotas.
Negotiations for the introduction of explicitly distributed
quotas commenced in December 1973 and were concluded
in 1977. There were four quota regulations, two of which
never took effect. All TACs adopted were higher than those
recommended as conflict over quotas caused the TAC to be
inflated to levels higher than initially agreed upon. The first
TAC set in 1974 was so high that harvests did not reach it.
However, recommendations for lower TACs were still
rejected and by the time a lower TAC was finally imposed,
it was too late. From 1977, directed fishing for herring in
the North Sea was banned.
Demise of the NEAFC
By the end of 1976 it was clear that the existing voluntary
regime was ineffective. The NEAFC was discredited by its
failure to adopt necessary and timely conservation measures
and by its inability to take necessary enforcement measures
(Mason, 1979). Negotiations over the 1977. quota
allocations broke down.
Following the proclamation of 200nm EEZs by most States
in 1977, all North Sea fisheries became subject to the
jurisdiction of coastal States. A new convention was
proposed, as some kind of multilateral forum was
considered desirable, but it was only to apply outside the
EEZs and to generate consultation and information
exchange rather than to manage stocks. In 1980 the NEAFC
was resurrected to fulfil this role under the auspices of a
new convention. However, no common fisheries
management system has yet been put in place (Oceans
Institute of Canada, 1990).
532
Although the situation would have been worse had it not
existed, the NEAFC nevertheless presided over a decline in
fish stock. It is suggested that by 1976 many stocks were
further from a state of ‘rational exploitation’ than they were
when the treaty was signed in 1959 (Underdal, 1980).
Northwest Atlantic Fisheries Organization
Administration
The Northwest Atlantic Fisheries Organization (NAFO) was
formed in 1979 under the Convention on Future Multilateral
Cooperation in the Northwest Atlantic Fisheries. The
Regulatory Area under the convention covers only the high
seas. NAFO was preceded by the International Fisheries
Commission for the Northwest Atlantic (ICNAF), which
was responsible for management of common fisheries prior
to the declaration of EEZs in the north-west Atlantic.
NAFO is constituted by a General Council, a Scientific
Council, a Fisheries Commission and a Secretariat. The
Commission makes proposals for joint action by the Parties
to the Convention designed to achieve the optimal utilisation
of fishery resources. These measures are also to promote
coordination and consistency between coastal State
conservation measures in the EEZ and those taken on the
high seas. The proposals are transmitted to the Parties and
become binding upon Parties which do not file an objection
to the proposal. That is, acceptance of management
measures is voluntary.
Management measures
NAFO utilises a range of conservation measures based upon
the notion of optimum yield, which is approximately 10%
more conservative as a mechanism for fisheries management
than maximum sustainable yield. Much of its fisheries
managementexpertise was inherited from ICNAF, including
considerable knowledge of the behaviour and status of
stocks. Measures adopted included minimum mesh sizes for
nets, closed areas and seasons, gear and vessel size
restrictions, minimum fish size limits, TACs and national
quotas for each principal commercial stock.
NAFO’s conservation measures in its Regulatory Area were
initially successful in the early 1980s and stocks of cod and
plaice showed signs of recovery. However, the
improvement was short-lived and they and other stocks have
since declined or remained at low abundance. The Scientific
Council has expressed concern regarding all stocks managed
by NAFO on the basis of scientific findings and reduced
catch per unit of effort. Cod, American Plaice, Redfish and
Yellowtail Flounder are each displaying weak recruitment
under fishing pressure and 1989 quotas on major stocks
were reduced to approximately two-thirds of those of 1988.
In recent years it has become apparent that NAFO has been
ineffective in its conservation efforts (Oceans Institute of
Canada, 1990).
The NAFO Joint Inspection Scheme allows Parties
reciprocal rights to board and inspect vessels. However,
there is no joint enforcement scheme. Breaches of the
regulatory measures must therefore be conducted by the flag
State.
Management crisis
The principal reasons for the recent failure of NAFO to
conserve fish stocks are disunity among the Parties and
continued fishing in the Regulatory Area by non-Parties.
Canada, for example, is reconsidering its participation in
NAFO.
Conflicts are continuing to take place between Canada and
the EC, in particular with Spain and Portugal. Spain has
been exceeding its quota for cod in the area designated as
3NO, where most cod is fished. Following the accession to
the EC in 1985 of Spain and Portugal, the EC objected to
TACs and quotas set in 1986, 1987 and 1988 for several
stocks, and increased its fishing effort for them. It has used
the objection procedure under the Convention to exempt
itself from various conservation measures. In 1989, it set
itself a quota more than 10 times that allotted to it by
NAFO. While the EC has been pressing for TACs to be
based upon maximum sustainable yield instead of optimum
yield and against protection measures for certain stocks of
cod, Canada wishes to conserve the juveniles and spawning
grounds of stocks which straddle its EEZ and the NAFO
Regulatory Area.
A range of countries fish in the Regulatory Area without
any formal commitment to conservation of its resources.
These include the USA, some Central and South American
countries, Republic of Korea and also EC vessels operating
under flags of convenience. Their catches are estimated to
exceed any surplus available following the allocation of
quotas to NAFO Parties.
North American Fisheries Commissions
International Pacific Halibut Commission
In 1923 Canada and the USA formed the International
Pacific Halibut Commission to restore Pacific Halibut
533
Management of International Resources
stocks by means of imposition of closed seasons. The
Commission’s conservation programme was revised in
1930, introducing an annual quota for each of four
management areas and a minimum size restriction on fish
landed. These measures were effective to increase stocks by
50% between 1932 and 1954. Supplementary arrangements
were entered into with Japan and the former USSR during
this period.
Although the quota system conserved the Pacific Halibut
stock, its economic effects were in some ways detrimental.
These included increased investment in gear and equipment
and increased fleet capacity. Fish tended to reach the
consumer in poorer condition, having been harvested earlier
in the season as a result of the rush to fill the quota.
The declaration of EEZs by Canada and the USA in 1976
brought the Pacific Halibut under national jurisdiction.
However, the Commission remains an example of
successful international management of a shared resource.
The fact that it was a bilateral rather than a multilateral
organisation signals the advantages of fewer participating
members. It also provides an example of effective use of
the quota system, although an incidental decline in quality
of fish reaching the consumer resulted.
USA administration
The USA has eight regional councils established under the
US Magnusson Fisheries Conservation and Management
Act. Most have adopted a species-by-species management
plan. The Act restricts the use of limited access as a
management tool, and rules out taxes or fees for domestic
fisheries. The USA also has three marine fishery
commissions for its Atlantic, Pacific and Gulf coastal states.
These were formed in the 1940s to coordinate the fisheries
of US states. Fish which are fully exploited and which are
harvested primarily in federally controlled waters (the EEZ)
must be brought under a Fisheries Management Plan.
Following proclamation of the 200nm EEZ in 1976, there
was a huge increase in the number of US fishing vessels
and this increased capacity led to the usual problems of
overfishing, decreasing incomes and so forth (Brown and
Crutchfield, 1981).
Alternative approaches
Harmonisation
States with common interests in fisheries management may
choose to harmonise their fisheries laws. This has been the
case in the south-west Pacific.
The South Pacific Forum Fisheries Agency (FFA),
established in 1979 as an arm of the South Pacific Forum,
has 13 members from the south Pacific region. In 1981 it
formed the Agreement Concerning Co-operation in
Management of Fisheries of Common Interest, which seeks
to coordinate regional fisheries policies and to harmonise
the management of fisheries, especially in the case of
common stocks. To this end, it standardises licensing
procedures, terms and conditions, and coordinates
surveillance and enforcement functions. Although there is
no limit on fishing effort, all fishing access agreements in
the region must comply with a harmonised list of access
3. Conservation and Management of Biodiversity
conditions and a regional register of licensed fishing vessels
is kept by participating local States.
In 1989, a Convention for the Prohibition of Fishing with
Long Driftnets in the South Pacific was concluded to
further harmonise fisheries management laws in the region.
It is administered by the FFA. The convention prohibits the
use of driftnets exceeding 2.5km in length by people or
vessels under the jurisdiction of the Parties within the
convention area, which includes both EEZs and the high
seas. Parties are to take action against any fishing using
driftnets in the Area by non-Parties, including prohibition
of landing fish caught by driftnets in their territory.
Property rights
Traditional open access to fisheries has permitted fishermen
ownership of the fishery resource on the basis of fishing
effort. Fisheries commissions limit the fishing effort. An
alteration of the property right may take the form of a
licence to fish, within an individual quota, which can be
freely traded and is itself an asset, without which the fish
cannot be owned.
It has been argued that the move to a system based on full
ownership with profit incentives would increase productivity
and efficiency, while at the same time removing the
"imperatives of the commons" (Keen, 1988). That is, the
rush to exploit the resource before others would be replaced
by an owner’s incentive to look after the property over the
longer term. Therefore, no imposition of limits upon fishing
effort would be necessary.
However, this approach relies upon the licensing of only so
many vessels as are required to harvest the resource. It
creates a windfall for those boat-owners permitted to remain
in an existing fishery and problems arise in identifying who
should be given the right to remain (Keen, 1988). It is also
suggested that such a scheme does not improve management
of the resource because fishermen will continue to increase
their harvesting capacity and compete to harvest the
resource. To be successful, therefore, fishing effort must be
effectively controlled, so that each fishing unit contains an
optimal combination of vessels, gear and so forth.
Ultimately, an independent regulatory body continues to be
necessary to oversee the process of licensing and effort
limitation.
Conclusions
Despite the declaration of EEZs and the introduction of new
management concepts, international fisheries commissions
remain necessary for the proper management of
international fisheries resources. This is because fish stocks
regularly cross international boundaries and fishermen
habitually compete to catch them.
However, most fisheries commissions have proved to be
relatively ineffective in the management of fisheries within
their competence. There are several reasons for this which
can be learnt from the operation of the commissions
detailed above. These concern the nature of the decision-
making processes involved and the mix of regulatory
measures used.
e Voluntary reporting does not provide reliable
information about fishery stocks; so independentresearch
is required. This may be provided to the commission
from a range of sources, including Member States.
e The process of deciding the amount of the TAC must be
kept separate from decisions on the allocation of quotas.
The primary decision concerning the TAC needs to be
made by a scientific committee and based on biological
rather than economic grounds.
e Distribution of quotas between States is best managed
either by a commission with few members or in a
situation where it is possible for States to engage in
bargaining for an exchange of various benefits brokered
by the commission.
e The quantitative approach to regulation is wasteful
because it regulates a catch after it has been caught.
Simpler and more enforceable restrictions should form
the basis of a regulatory system. An appropriate mix of
measures would centre around gear, area, season and
duration restrictions, which are more amenable to
enforcement within port.
e The option of a unilateral objection procedure
undermines the delicate compromises which a resource
distribution involves. A commission needs a strong
central authority to overcome disagreements between its
members.
e Enforcement of international management decisions by
Member States against their own nationals tends to be
lax. A commission needs direct enforcement powers
against recalcitrant members and fishermen in order to
ensure that its recommendations are put into practice.
e Licensing and radar surveillance are more economical
and efficient systems of monitoring»than the current
system of inspections. Reciprocal observation
arrangements, such as employed by the NAFO can
supplement such a system.
The uninspiring performance of fisheries organisations to
date need not be taken as conclusive of their ineffectiveness.
Where the members of an international commission have
the will to conserve fisheries cooperatively, measures can
be designed to implement effective fisheries conservation.
ANTARCTICA: THE EVOLUTION OF AN
INTERNATIONAL RESOURCE MANAGEMENT
REGIME
Geography
The Antarctic region, which includes the Southern Ocean as
well as the continent itself and its islands, is the largest
wilderness left in the world (Laws, 1989). The region
covers 13.918 million km?, which is almost 10% of the
earth’s surface.
The continent is the driest, highest and coldest in the world,
and is almost entirely covered by ice. During winter the sea
ice rapidly increases round the continent and adds a further
20 million km? to the size of the ice cap. In some places the
ice cap is estimated to be as much as 4.7km thick.
The Southern Oceans are some of the most turbulent in the
world (Techernia and Jeannin, 1983). There are two main
currents. Close to the coast a westerly current predominates
while further from shore the main current is easterly; the
interface between these is the Antarctic Divergence, a
complex shear zone of upwelling where nutrient-rich deep
water is brought to the surface, thereby providing the
primary basis of the Southern Ocean food web (Deacon,
1987). Another important feature of this ocean is the
Antarctic Convergence, where cold surface waters plunge
beneath the warmer and less dense subtropical waters at
around 50°S. The exact location of this convergence is not
fixed but the pronounced changes in temperature and
salinity on either side are relatively constant (Holdgate,
1984).
First human contact
The first recorded human contacts with the region were
during Cook’s voyages into the Southern Ocean between
1772 and 1775 (Beaglehole, 1961). Shortly after this, the
huge populations of seals attracted sealers to the region,
thus initiating the cycle of over-exploitation, collapse and
regeneration typical of open-access resources (Bonner,
1968). Sealers at first concentrated on the islands and by
1822 (Bonner, 1968) many populations had collapsed; an
estimated 1.2 million fur seals had been taken in South
Georgia and one million in the South Shetland region.
Sealers remained active in the region for the next 100 years
and seal numbers did not recover to estimated pre-
exploitation levels until recent decades (Bonner, 1982).
The Southern Oceans support many cetaceans, including the
large and commercially valuable Blue, Fin and Sei Whales.
Whalers lacked the technology to capture and process these
whales until the 1870s, but once these difficulties were
overcome, they moved into the region and operated for
several decades from shore-based processing facilities on
sub-Antarctic islands. Factory ships were first used in the
1925-26 season; these allowed whales to be processed at sea
thereby greatly increasing the number caught (Bonner,
1980). Efforts were initially concentrated on the species
with highest commercial value, such as Humpbacks, Blue,
Fin and Sei Whales, but as the stocks of these declined,
attention turned to other species (Bonner, 1984). By the
latter half of this century only the Minke, smallest of the
more common baleen whales, had not been subject to
intensive commercial harvesting (Bonner, 1980).
Sovereign States of Antarctica
Seven States have made claims of territorial sovereignty in
Antarctica which they have defined. These claims are based
on a variety of doctrines such as: discovery, formal
annexation, sector theory and occupation (Kish, 1973). In
addition, both the USA and the USSR have maintained that
they have a basis for such claims, although they have not
made specific claims themselves and do not recognise the
claims of the other States. Even though the validity of these
claims may be dubious under modern principles of
international law (Greig, 1988), it should be recognised that
for the claimant States they are made seriously and some
States may not care to relinquish them (Conforti, 1986).
The whaling treaties
The first international resource management commission
535
Management of International Resources
which included the region within its jurisdiction was the
commission established under the 1931 Convention for the
Regulation of Whaling. This convention prohibited
commercial whaling of two depleted species, Right Whale
and Bow Whale, and banned the killing of calves and
immature or female whales in the company of calves or
sucklings. It further required whalers to make full use of
the carcasses. However, the convention had little practical
effect as several major whaling nations refused to accede to
it.
The successor to this convention, the International
Convention for the Regulation of Whaling, came into force
in 1948 and was ratified by most major whaling nations. Its
basic aim was to control whaling so as to avoid over-
exploitation and to ensure conservation of the stocks
(Birnie, 1985; Rosati, 1984). The convention established the
International Whaling Commission (IWC) to implement the
aims of the convention and regulate whaling by establishing
quotas and acceptable methods of capture, and designating
protected species (Smith, 1984). The schedule to the
convention contains regulations governing the protection
and exploitation of whales, listing protected species and
setting quotas for others. The schedule may be amended by
a three-quarter majority of the members at the annual
general meeting of the IWC, which is composed of
representatives of each contracting party to the convention.
Initially, the IWC set annual quotas based on the "Blue
Whale unit", which essentially meant that the whaler could
take any combination of whales of any species up to the
equivalent mass of the number of Blue Whales that had
been allocated. As a result, whales which were more
valuable per unit weight were more heavily exploited until
their stock numbers had collapsed, whereupon the next most
valuable stock was exploited.
Under this regime the industry continued to grow, and
numbers taken worldwide increased year after year,
reaching a peak in the 1960/1961 season when
approximately 64,000 whales were killed (International
Whaling Commission, 1963). This mechanism of setting
quotas, which proved to do little to conserve whales, was
abandoned in 1972 in favour of quotas on a species basis.
This new approach was then later enhanced by the "New
Management Procedure" which established quotas based on
a stock-by-stock approach (Birnie, 1982). This tightening in
procedure was also accompanied by a reduction in the
number of whales which were allowed to be harvested in
any year (Birnie, 1989). As a result of this and other
factors, catches declined steadily until 1982 when the IWC
declared a worldwide ‘pause’ on commercial whaling
effective from the 1985/1986 season, which is still in force.
The Antarctic Treaty
The Antarctic Treaty (‘the Treaty’) was adopted in 1959
and came into force in 1961. It is essentially a self-denying
ordinance under which contracting parties agree: to prevent
military activity in the area and to use Antarctica for
peaceful purposes only; to promote international cooperation
in scientific research; and to ban nuclear explosions and
disposal of radioactive waste. Also within its articles, the
Treaty preserves all existing rights and claims to
3. Conservation and Management of Biodiversity
sovereignty and the position of those who recognise no
claims, and nullifies any basis of claim during its operation.
The Treaty is a classic example of a ‘framework
convention’, with the Treaty itself being quite short and
general in nature, leaving matters of detail to be negotiated
at a future time through recommendations or protocols. In
addition, the Treaty established no independent institutional
structure (Secretariat) for its implementation. Membership
of the Treaty is open to all countries, with two categories
of members, Consultative Parties and Non-Consultative
Parties. In order to become a Consultative Party a country
has to display a serious interest in Antarctica as
demonstrated by substantial scientific research activity in
the region. Consultative Parties have voting rights in the
Antarctic Treaty system and, therefore, are responsible for
the governance of the region, while Non-Consultative
Parties merely have observer status at the meetings of the
Parties. Details of the present membership of the Treaty are
given in Table 33.3.
Periodic meetings of the Parties are held to exchange
information, consult on matters of common interest
pertaining to Antarctica, and formulate measures to manage
and govern the region. These meetings, called Antarctic
Treaty Consultative Meetings (ATCM) (Myhre, 1986),
occur every two years at a conference hosted and organised
by one of the Consultative Parties. In addition, special
meetings (SATCM) are called from time to time to consider
specific issues. Recommendations are made at meetings on
a consensual basis. To date, nearly 200 recommendations
have been made on a wide variety of subjects, including:
protection of the environment, meteorology,
telecommunications, transport and logistics, tourism and
exchange of information.
In the Treaty itself there is only one brief but
comprehensive reference to environmental matters: a short
provision calling upon the Consultative Parties to develop
measures for the "preservation and conservation of the
living resources of Antarctica". From this general
obligation, an elaborate management regime has been
developed through additional recommendations, protocols
and further conventions to provide comprehensive
protection for the environment in Antarctica.
Agreed Measures for the Conservation of Antarctic
Fauna and Flora
The first significant development pursuant to the general
obligation to protect and conserve the Antarctic environment
occurred in 1964 with the adoption of the Agreed Measures
for the Conservation of Antarctic Fauna and Flora (the
Agreed Measures) which represent one of the earliest
examples of effective international regulation of a resource.
The general intention behind this recommendation was to
protect the living resources of Antarctica. In particular, it
covered protection of mammal and bird life from
unnecessary slaughter, and the minimisation of disturbance
on land by personnel from the growing number of scientific
bases.
Over and above the general provisions (which are applicable
to the entire Treaty Area), the Agreed Measures allow for
more stringent provisions with regard to ‘Specially
Protected Areas’ (SPA) and ‘Specially Protected Species’.
536
Permits may only be granted by a contracting party for the
taking of Specially Protected Species "for compelling
scientific purposes” and even then they may not "jeopardise
the existing natural ecosystem or survival of that species".
Areas which are designated as Specially Protected Areas are
protected by similarly stringent provisions (Anderson,
1968).
Criteria for review of the SPAs and for establishing Sites of
Special Scientific Interest (SSSI) to protect sites important
for research were developed later. However, the areas
designated as either SPA or SSSI have been relatively small
in size. A recent review by the Scientific Committee on
Antarctic Research (SCAR), which acts as the scientific
committee for the Agreed Measures, considered the existing
SPAs and SSSIs and concluded that some areas worthy of
designation remained undesignated and that the
documentation of individual sites remained uneven and
incomplete.
During the late 1980s moves were made to remedy these
defects and, as a result, two further categories were
established. The Specially Reserved Area is intended to
protect representative examples of major geological features
and those of outstanding aesthetic, scenic and wilderness
value while the Multiple Use Planning Area (MPA) is a
mechanism for controlling human activities in high-use
areas, to minimise harmful environmental impacts.
The protected areas system has been rationalised under the
provisions of Annex V to the Madrid Protocol, which
introduced the Antarctic Specially Protected Area (ASPA)
and the Antarctic Specially Managed Area (ASMA). In due
course existing SPAs and SSSIs will be re-designated as
ASPAs, and MPAs will be re-designated as ASMAs.
Currently there are 19 SPAs, 35 SSSIs, 1 Specially
Reserved Area and 1 Multiple Use Planning Area (see Fig.
33.1 and Table 33.4).
The Convention for the Conservation of Antarctic Seals
Since the 1780s, seals had been subject to gross over-
exploitation, with population collapses occurring in the
1820s and again in the 1860s. By the 1960s, seal numbers
and stocks were returning to their pre-exploitation levels
(Mitchell and Tinker, 1980), when Norway expressed
interest in recommencing commercial exploitation.
Commercial sealing has not in fact been re-established;
nevertheless, Norway’s actions caused considerable concern
and moves were made to bring about a legal instrument to
control exploitation of seals.
The conference of the Parties to the Antarctic Treaty once
again provided the forum within which- resource
management could occur. In 1964 it was suggested that
national governments should regulate pelagic sealing on a
voluntary basis (Myhre, 1986), followed two years later by
the adoption of Interim Guidelines for the Voluntary
Regulations of Antarctic Pelagic Sealing (Recommendation
IV-XX]). Finally, in 1972 Consultative Parties adopted the
Convention for the Conservation of Antarctic Seals, which
came into force in 1978 (Lyster, 1985) and is renewed
every five years.
Management of International Resources
Table 33.3 Parties to the Antarctic Treaty
CONTRACTING PARTIES {in chronological order)
United Kingdom*
South Africa*
Belgium*
Japan*
USA*
Norway*
France*
New Zealand*
Commonwealth of Independent States *®
Poland
Argentina*
Australia*
Chile*
Czechoslovakia
Denmark
Netherlands
Romania
Germany, DDR#
Brazil
Bulgaria
Germany, BRD#
Uruguay
Papua New Guineae
Italy
Peru
Spain
China
India
Hungary
Sweden
Finland
Cuba
Korea, Rep
Greece
Korea, Dem. People’s Rep
Austria
Ecuador
Canada
Colombia
Switzerland
Guatemala
t++etteeetgeeetet
++ ttt
31 May 1960 1
21 June 1960 2
26 July 1960 3
4 August 1960 4
18 August 1960 5
24 August 1960 6
16 September 1960 7/
1 November 1960 8
2 November 1960 9
8 June 1961 (29 July 1977) 10
23 June 1961 11
23 June 1961 12
23 June 1961 13
14 June 1962 14
20 May 1965 15
30 March 1967 (19 November 1990) 16
15 September 1971 17
19 November 1974 (5 October 1987) 18
16 May 1975 (12 September 1983) 19
11 September 1978 20
5 February 1979 (3 March 1981) 21
11 January 1980 (7 October 1985) 22
16 March 1981 23
18 March 1981 (5 October 1987) 24
10 April 1981 (9 October 1989) 25
31 March 1982 (21 September 1988 26
8 June 1983 (7 October 1985) 27
19 August 1983 (12 September 1983) 28
27 January 1984 29
24 April 1984 (21 September 1988) 30
15 May 1984 (9 October 1989) 31
16 August 1984 32
28 November 1986 (9 October 1989) 33
8 January 1987 34
21 January 1987 35
25 August 1987 36
15 September 1987 (19 November 1990) 37
4 May 1988 38
31 January 1989 39
15 November 1990 40
31 July 1991 41
Source: Scott Polar Research Institute, Cambridge.
Note: Made 1 December 1959; came into force 23 June 1961. The Treaty has no limits on its duration. It may be reviewed, at the request of the
Consultative Party.
Key: * Original signatories; the 12 states which signed the Treaty on 1 December 1959; the dates given are those of the Deposition of the
instruments of ratification, approval, or acceptance of the Treaty. + Consultative Parties; 26 states, the 12 original signatories and 14 (formerly
15#) others which achieved this status after becoming actively involved in Antarctic research (with dates in brackets). e Papua New Guinea succeeded
to the Treaty after becoming independent of Australia. # The two German states unified on 3 October 1990. Thus there are now 40 member states
from the 41 adherents. ® Comprising 11 of the reassociated 15 Republics of the Soviet Union, December 1991.
The object of this convention is to "promote and achieve the
objects of protection, scientific study and rational use of
Antarctic Seals and to maintain a satisfactory balance with
the ecological system". The convention covers six species
of seal: Southern Elephant Seal, Leopard Seal, Weddell
Seal, Crabeater Seal, Ross Seal and Southern Fur Seal.
Like the Agreed Measures, this convention operates by a
system of permits which allows the capture or killing of
seals for certain purposes and, in an Annex, establishes
quotas for commercial sealing. The measures outlined in the
Annex also seek to control other aspects of sealing through
537
establishing a sealing season, the areas in which sealing
may take place, and the methods which may be used to
capture and kill seals. In order to monitor properly the
taking of seals, the convention also requires the Parties to
report to SCAR all seals which have been taken in any one
season and to report annually on the steps that they have
taken to implement the convention. The convention also
provides for the establishment of a Commission and
Scientific Advisory Committee if and when commercial
sealing is re-established in the region.
3. Conservation and Management of Biodiversity
Figure 33.1 Protected areas in Antarctica
009 OOF O02
C= a
Lu»
uolydefoi1g s1ydDIibOe8198}S IDjOY
538
Table 33.4 Protected areas in Antarctica
AREA DATE
(ha) ESTAB.
Specially Reserved Area
1 North Dufek Massif 48,000 1991
Specially Protected Area
2 Ardery Island and Odbert Island 220 1966
3 Avian Island, North-West Marguerite Bay 40 1990
4 Beaufort Island 1,865 1966
5 Cape Hallett 25 1966
6 Coppermine Peninsula 65 1970
7 Cryptogam Ridge, Mount Melbourne 60 1991
8 Dion Islands 100 1966
9 Forlidas Pond and Davis Valley ponds 600 1991
10 Green Island 25 1966
11 Lagotellerie Island 130 1985
12 Litchfield Island 250 1975
13 Lynch Island 10 1966
14 Moe Island 100 1966
15 New College Valley 10 1985
16 North Coronation Island 5,000 1985
17 Rookery Islands 65 1966
18 Sabrina Island 60 1966
19 Southern Powell & adjacent islands 610 1966
20 Taylor Rookery 30 1966
Site of Special Scientific Interest
21 Ablation Point - 18,000 1990
Ganymede Heights, Alexander Is
22 Ardley Island 300 1991
23 Arrival Heights 60 1975
24 Barwick Valley 29,120 1975
25 Biscoe Point 200 1985
26 Byers Peninsula 3,027 1975
27 Canada Glacier 100 1985
Management of International Resources
AREA DATE
(ha) ESTAB
28 Cape Crozier 462 1975
29 Cape Royds 2 1975
30 Cape Shirreff 265 1989
31 Caughley Beach 25 1985
32 Chile Bay (Discovery Bay) 75 1987
33 Cierva Point 850 1985
34 Clark Peninsula 800 1985
35 East Dallman Bay 60,000 1991
36 Fildes Peninsula 154 1975
37 Harmony Point 1300 1985
38 Haswell Island 80 1975
39 Linnaeus Terrace 300 1985
40 Lions Rump 100 1991
41 Marine Plain, Mule Peninsula 2,340 1987
42 Mount Flora, Hope Bay, Antarctic Peninsula 65 1990
43 North-east Bailey Peninsula 100 1985
44 North-west White Island 1350 1985
45 Parts of Deception Island 100 1985
46 Port Foster, Deception Island 50 1987
47 Potter Peninsula 200 1985
48 Rothera Point 4 1985
49 South Bay, Doumer Island 70 1987
50 Summit of Mt Melbourne 800 1987
51 Svarthamaren 390 1987
52 Tramway Ridge 1 1985
53 Western Bransfield Strait 103,000 1991
54 Western Shore, Admiralty Bay 160,000 1979
55 Yukidori Valley 300 1987
Multiple Use Planning Area
56 South-west Anvers Island 153,500 1991
Source: Swithinbank, C. 1991. Conservation areas of Antarctica. Unpublished contribution to the Antarctica Digital Database, in preparation.
WCMC/BAS/SPRI, Cambridge.
The Convention on the Conservation of Antarctic
Marine Living Resources (CCAMLR)
During the 1960s Japan and the USSR began investigating
the possibility of commercial harvesting of krill, a shrimp-
like crustacean. As krill play a central and vital role in the
entire region’s ecosystem (Auburn, 1982) the Consultative
Parties decided to establish a legal regime to control this
emerging industry before it developed (Barnes, 1982).
Once again the Antarctica Treaty System provided the
framework and negotiations began in 1977, followed by
seven official meetings and consultations on the proposed
convention (Edwards and Heap, 1980). The two central
concerns that shaped the resulting convention were the role
played by krill in the food chain in the Antarctic region and
the ever-present conflict of the territorial claims.
Krill has been the driving force behind the parties’
recognition of the need to consider the Antarctic ecosystem
in total. This is because krill is a key factor in the food
chain in the Antarctic region; it feeds on plankton and, in
turn, is the principal food for many species of birds, fish
and whales (Mitchell and Sandbrook, 1980). Because the
food chain in the Antarctic is simplified, with relatively few
species, the mass removal of one species which is central to
it is likely to have very significant effects on the whole
539
chain. Consequently, should there be any serious
disturbance to the krill, this will have serious implications
for the entire Antarctic ecosystem. A further consideration
is that krill has a tendency to swarm into large, densely
packed formations, making it relatively easy to harvest on
a large scale and particularly susceptible to over-
exploitation. Therefore, even though the ostensible purpose
of the negotiations was to protect krill from over-
exploitation, the main concern was for the effects that
overharvesting would have on other species in the
Antarctic. This concern is reflected in CCAMLR in the
“ecosystem approach’ adopted by the convention (Edwards
and Heap, 1980).
The other issue which dominated negotiations was the legal
debate arising from the conflicting territorial claims.
Although these had been frozen by the Treaty in 1959, by
the late 1970s the potential wealth these claims represented
was closer to becoming reality; so claimants were anxious
to ensure that nothing in CCAMLR would prejudice their
claims or rights to exercise jurisdiction over the coastal
waters. Similarly, the non-claimant States were anxious to
ensure that CCAMLR did not legitimise or help legitimise
the position of the claimants (Triggs, 1987).
These issues are representative of the constraints to
effective regulation of international resources. Ecosystems
generally overlap state boundaries, which leads to need for
3. Conservation and Management of Biodiversity
international environmental cooperation. It is the States’
refusal to relax claims to absolute sovereignty, even in the
face of obvious gains from cooperation, which leads to the
difficulties of achieving effective international
environmental regulation. The Antarctic system has been
virtually unique in its ability to cope with both sources of
conflict.
By September 1978 the key issues had been resolved and
were the subject of a ‘gentlemen’s agreement’, which lasted
until the convention came into force in 1982.
The object of CCAMLR is the "conservation of Antarctic
marine living resources". Because of the ecosystem
approach applied to achieve this object, the convention’s
application extends to all areas within the Antarctic
Convergence. The coverage of CCAMLR is, therefore,
considerably larger than the Treaty itself. This defining of
the area covered by the convention on biological grounds as
compared to political ones is an innovative feature of this
convention.
Within the area covered by the CCAMLR, however, there
are a number of islands which are the undisputed territory
of Consultative Parties, and which are not covered by the
convention. The existence of these islands and the fact that
they are outside the jurisdiction of CCAMLR is recognised
in Article [V(2)(b). This Article provides that "nothing in
this Convention and no acts or activities taking place while
the present Convention is in force shall: ... be interpreted
as a renunciation or diminution ... of, or as prejudicing,
any right or claim or basis of claim to exercise coastal state
jurisdiction under international law within the area to which
this Convention applies ...". This provision deliberately
does not refer to the undisputed islands, an ambiguity which
allows claimant States to interpret the provision as also
referring to the disputed claims within the Antarctic Treaty
area (south of 60°S), whereas the non-claimant States can
interpret the provision as meaning that CCAMLR applies
only to where national sovereignty is generally recognised.
This ambiguity therefore allows the Consultative Parties to
come to agreement on the issue of conserving the marine
living resources while apparently maintaining the status quo
on territorial claims.
The ecosystem approach adopted in this convention means
that it is unlike most other fishery agreements which set
quotas based upon maximum sustainable yields (Gulland,
1968; Bean, 1983). CCAMLR sets a standard based not
only on the maximum sustainable yield of the target species
but also requires that equal consideration be given to the
likely effects on other species and the marine ecosystem as
a whole.
CCAMLR, for the first time in the Antarctic Treaty system,
establishes a commission to implement its objectives, the
Commission for the Conservation of Antarctic Marine
Living Resources. The Commission has its headquarters in
Hobart, Australia, and is composed of delegates from all
Contracting Parties; it is the first permanent resource
management commission established for the continent.
CCAMLR also establishes a scientific body to act as a
consultative body to the Commission.
540
To ensure that the provisions of the Convention are
observed in the absence of any binding settlement procedure
for disputes, the Convention adopts a number of
conventional mechanisms. Each Contracting Party is
required to "take appropriate measures within its
competence to ensure compliance with the provisions of the
Convention and with conservation measures adopted by the
Convention...". To facilitate compliance with the
Convention further, CCAMLR also establishes an elaborate
system of observation and inspection. Contracting Parties
are also required to make extensive annual reports to the
Committee. A distinctive feature of the convention is the
obligation on the Commission to notify other Contracting
Parties of the infringements of the convention by any one
Contracting Party, thereby hoping to ensure observance
through public opprobrium.
Convention on the Regulation of Antarctic Mineral
Resource Activities (CRAMRA)
The Antarctic region is expected to contain enormous
mineral wealth but, despite numerous geological surveys,
no commercially viable deposits have been discovered as
yet. However, mindful of the enormous impact that a full-
scale mining operation would have on the sensitive
Antarctic environment, the Consultative Parties, after
CCAMLR finally entered into force in 1982, began serious
negotiations for a similar treaty for the regulation of the
development of the mineral resources (Rich, 1981). Prior
to this, a moratorium on mineral resource activity in
Antarctica had been declared in 1977, and subsequently
extended, dependent on the "timely conclusion cf a
convention on mineral resources activity".
CRAMRA was the most detailed and complex of the legal
instruments making up the Antarctic Treaty System. It
essentially aimed to create a regime where mineral resource
activity could not take place until the proponent of such an
activity could prove that the activity in question would not
cause damage to the Antarctic environment. The placing of
the onus of proof on the developers rather than the
regulators in this convention is a major advance.
CRAMRA also provided for the establishment of five new
resource management institutions. Implementation of the
Convention would have been overseen by the Antarctic
Minerals Resources Commission, which would have
consisted of the representatives of the Consultative Parties
and other nations engaged in or sponsoring mineral
research. They would have been advised by a Scientific,
Technical and Environmental Advisory Committee, and
Regulatory Committees would have been established for
each area identified by the Commission where resource
activity could take place. A Secretariat would have been
established to service these bodies and the special meetings
of the Contracting Parties convened to discuss mineral
resource issues.
In 1988, after six years of negotiation, CRAMRA was
finally adopted and opened for signature. However, after
considerable public lobbying, the Consultative Parties
decided not to ratify the convention. In May 1989 it was
declared that Australia would not ratify CRAMRA because
they believed that no mining should take place at all in the
region. Support for the Australian position came from the
French government, and in August 1989 the two
governments issued a statement to the effect that mining
was incompatible with protection of the Antarctic
environment and indicated that they would not ratify
CRAMRA but would pursue negotiation of a comprehensive
environment protection convention within the framework of
the Treaty. These two countries then submitted a joint
working paper proposing the preservation of Antarctica as
a ‘Wilderness Reserve’ (Antarctic Treaty Consultative
Meeting 1990a,b). Similarly, several other Consultative
Parties indicated that they would not ratify CRAMRA, with
some of them also submitting proposals for a
comprehensive protection regime (Redgwell, 1989).
Although there has been no formal recognition by the
parties that CRAMRA is defunct, it is generally accepted
that CRAMRA has been overtaken by events and is no
longer going to enter into force. Even so, many of the
techniques developed, such as the ‘onus of proof’ in the
environmental impact assessment being shifted from the
regulator to the developer or the extensive provisions on
institutional inspection to ensure compliance within the
convention, are of more than passing historical interest. For
not only do many of them reappear in CRAMRA’s
successor, the protocol for the comprehensive environment
protection regime, but they are of interest in the precedent
that they establish for future legislation.
The comprehensive environment protection regime
A Special Antarctic Treaty Consultative Meeting was called
to consider the various proposals submitted by the
Consultative Parties in relation to a comprehensive
environment protection regime. The first session of
SATCM was held in Chile in 1990; it agreed a draft
protocol to the Treaty which formed the basis of discussion
for a meeting in Madrid in 1991. At this second session a
new draft protocol was agreed and recommended to
governments for formal consideration. This draft protocol
was generally well received except for the clause dealing
with the amendment of the protocol. Eventually a
compromise was reached which effectively placed a 50-year
moratorium on mining in the region, after which time any
Party to the Treaty could request a review. Any proposal to
amend the moratorium can only become binding if it
receives the approval and acceptance of three-quarters of
the Consultative Parties, and only then if there is an agreed
binding legal regime to regulate mineral activities. If such
a modification has not entered into force within three years
of the date of its adoption, any Party may withdraw from
the protocol with two years’ notice. With the resolution of
this final problem the way was open for the ‘Madrid
Protocol’ to be adopted in Spain, just before the XVIth
ATCM in Bonn, Germany in October 1991.
The object of the protocol is to establish a "comprehensive
regime for the protection of the Antarctic environment and
dependent and associated ecosystems and hereby designate
Antarctica as a natural reserve, devoted to peace and
science". It seeks to build upon the Antarctic Treaty System
by consolidating the sometimes disparate elements of the
system to create a comprehensive regime. The protocol,
541
Management of International Resources
like the Treaty, is a framework within which general
obligations are agreed to, which will be translated into
specific procedures and management guidelines in future
annexes to the protocol. Features of this framework are:
e The environmental principles by which Parties should
plan their activities in Antarctica (Article 3);
The obligation for Parties to cooperate fully in the
planning and conduct of activities in the Treaty Area
(Article 6);
A general moratorium on mineral resource activity other
than for scientific research (Article 7);
The establishment of a Committee for Environmental
Protection to help Parties implement the aims of the
protocol (Articles 11 and 12);
The establishment of a system of inspections to monitor
the observance by the Parties of the protocol (Article
14);
The requirement for Parties to submit annual reports
detailing the steps they have taken to implement the
Protocol (Article 17);
The establishment of Dispute Resolution Procedures
(Article 18 and 19).
More detailed annexes have also been developed, covering
environmental impact assessment (Annex I), conservation of
flora and fauna (Annex II), waste disposal and management
(Annex III), prevention of marine pollution (Annex IV) and
area protection and management (Annex V), which
introduces the Antarctic Specially Protected Area and the
Antarctic Specially Managed Area. Under Article 9 of the
Madrid Protocol the annexes form an integral part of the
protocol itself, and provision is made for subsequent
additional annexes to be adopted at a later date.
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Kish, J. 1973. The Law of International Spaces.
Knight, H.G. (Ed.) 1975. The Future of International Fisheries
Management. West Publishing Co., Minn.
Laws, R.M. (Ed.) 1989. Antarctica: the last frontier. Boxtree,
London.
Lyster, S. 1985. International Wildlife Law. Grotius Publications Ltd.
p.48.
542
Mason, C.M. (Ed.) 1979. The Effective Management of Resources: the
international politics of the North Sea. Pinter.
Mitchell, B. and Sandbrook, 1980. The Management of Southern
Oceans. ED, London.
Mitchell, B. and Tinker, J.
Earthscan, London.
Myhre, J.D. 1986. The Antarctica Treaty System: politics, law and
diplomacy. Westview Press.
Oceans Institute of Canada 1990. Managing Fishery Resources Beyond
200 Miles: Canada’s options to protect Northwest Atlantic
straddling stocks. Report prepared for the Fisheries Council of
Canada.
Redgwell, C.J. 1989. Antarctica. (Current Developments: Public
International Law). International and Comparative Law Quarterly
39:474-481.
Rich, R. 1981. A minerals regime for Antarctica. International and
Comparative Law Quarterly 31:709-725.
Rosati, J.P. 1984. Enforcement questions of the International Whaling
Commission: are. exclusive economic zones the solution? California
Western International Law Journal 14:114-147.
Rose, G.L. 1991. Community Fisheries Management Legislation in EC
Waters. CIEL, London.
Smith, G.A. 1984. The International Whaling Commission: an analysis
of the past and reflections on the future. Natural Resources Law
16:543-567.
Techernia, P. and Jeannin, P.F. 1983. Quelques aspects de la
circulation oceanique Antarctique révélés par l’observation de la
dérive d’icebergs (1972-1983). CNRS, Muséum National
d’Histoire Naturelle.
Triggs, G.D. (Ed.) 1987. The Antarctica Treaty Regime: law,
environment and resources. Cambridge University Press.
Underdal, A. 1980. The Politics of International Fisheries
Management. Oslo.
1980. Antarctica and its Resources.
Authors as follows: International fisheries, Greg Rose
(CIEL); Antarctica, Sam Johnston.
Current Practices in Conservation
34, CURRENT PRACTICES IN CONSERVATION
Chapters 28 to 33 have outlined the major instruments and
mechanisms, both national and international, which are or
can be used in the conservation of biodiversity. This chapter
examines some of the most important ways in which
conservation is carried out.
Actions to maintain biodiversity can be focused on three
levels: ecosystem diversity, species diversity, and genetic
diversity. All three are inextricably interlinked, but are not
synonymous: maintenance of ecosystem diversity implies
maintenance of the species (or at least the most important
species) which constitute that ecosystem; however, it is
perfectly feasible to maintain species independent of the
ecosystems or habitats in which they normally occur.
Similarly, maintenance of genetic diversity within a species
self-evidently implies maintenance of that species, although
the reverse does not apply, in that species can generally be
maintained at far lower levels of genetic diversity than
would be expected to occur under natural conditions.
However, at whatever level the problem is looked at, it is
axiomatic that the maintenance of species diversity, and in
particular the prevention of species extinctions, is pivotal to
the conservation of biodiversity.
The preservation of species as part of a functioning,
although not necessarily pristine, ecosystem is regarded as
in situ conservation. This is, and will remain, by far the
most important form of biodiversity conservation, for a
variety of reasons which will be discussed below.
Maintenance of species away from their normal habitat is
termed ex situ or off-site conservation. The distinction
between these forms of conservation is not absolute and
becomes increasingly blurred as individual species are made
the subject of complex, interventionist management
strategies.
Planning for the conservation of diversity can be
approached in two ways: habitat- or ecosystem-based and
species-based.
Habitat or ecosystem approaches
An ecosystem approach to conservation attempts to ensure
that representative samples of ecosystems or important
habitat types are maintained, through the designation of a
network of protected areas or through other controls on
land-use. It is assumed that by so doing, the species which
inhabit these ecosystems will be conserved. The principal
advantage of such an approach is that it does not require
detailed knowledge of the status and distribution of all
species, i.e. it can be assumed to protect species for which
information is not available. This applies particularly to
tropical rain forests, whose diversity is at present
unquantifiable because it consists largely of undescribed
species. A significant proportion of these are expected to be
given some measure of protection merely by the protection
of large areas of habitat. Its major drawbacks are the
difficulty of devising satisfactory habitat or ecosystem
classifications on which to base protected area networks,
and that fact that populations of particularly rare
543
and threatened species (i.e. those in most urgent need of
conservation) are likely in many cases not to be included in
a network of protected areas set up on the basis of
representative samples of major ecosystem types.
Species-based approaches
Species-based approaches entail the review of taxa with the
aim of identifying species considered to be of high priority
for conservation, most importantly threatened species and
those of actual or potential resource value. Conservation or
recovery plans can then be developed for these species,
often entailing a combination of in situ and ex situ
management. This approach is exemplified internationally
by the IUCN Red Data Books (Table 34.1), which treat the
status and conservation requirements of globally threatened
species in detail, and the work of the IUCN Species
Survival Commission (SSC).
Since its small beginnings in 1949, the SSC has grown into
a large global network. In 1991 it consisted of some 95
Specialist Groups with approximately 3,500 members in 135
countries. Through its members and the work of its
Specialist Groups, SSC promotes action to arrest the loss of
the world’s biological diversity and to restore threatened
species to safe and productive population levels. The SSC
is divided into Specialist Groups organised primarily on a
geographical and/or taxonomic basis, although there are
some ‘interdisciplinary’ groups. Among the existing taxon-
based Groups are: Antelopes, Parrots, European Reptiles
and Amphibians, Coral Reef Fish, Ants, Cycads,
Carnivorous Plants, Orchids; while examples of
interdisciplinary groups include Re-introductions and
Ethnozoology. Membership of Specialist Groups is purely
voluntary, and consists mainly of scientists and
conservationists nominated by the group Chairmen, who are
in turn appointed by the SSC Chairman. All appointments
are ratified by Council. Some of the larger Groups (e.g.
Captive Breeding) have established secretariats and employ
paid staff to accomplish their core activities.
The preparation of ‘Action Plans’ is one of the most
important activities undertaken by the SSC groups. Under
the current Action Planning programme, which started in
1986, each taxon-based Specialist Group is expected to
review the conservation status and needs of the species
within its remit, and recommend conservation actions which
will ensure their long-term survival. These
recommendations may include both in situ measures, such
as the carrying out of population surveys, gazetting of
particular sites as protected areas; provision of funds or
equipment to local enforcement agencies etc., and ex situ
measures, such as the establishment of captive breeding
populations. By early 1992, Action Plans covering 16
groups had been published by IUCN (Table 34.2), and
many more were in preparation.
Once Action Plans are published, the Specialist Groups
have a duty to promote the implementation of their
recommendations by lobbying governments, conservation
organisations and donors.
3. Conservation and Management of Biodiversity
Table 34.1 IUCN Red Data Books
GROUP YEAR
MAMMALS Mammal Red Data Book: the Americas and Australasia (excluding Cetacea) 1982
Threatened Primates of Africa 1988
Lemurs of Madagascar and the Comoros 1990
Dolphins, Porpoises and Whales of the World 1991
BIRDS Threatened Birds of Africa and Related Islands* 1985
REPTILES Amphibia-Reptilia Red Data Book: Testudines, Crocodylia, Rhynchocephalia 1982
INVERTEBRATES Invertebrate Red Data Book 1983
Threatened Swallowtails of the World 1985
PLANTS Plant Red Data Book 1978
Note: * An ICBP/IUCN Red Data Book. ICBP = International Council for Bird Preservation. See references for full citations.
Table 34.2 IUCN/SSC Action Plans
GROUP YEAR
MAMMALS African Insectivora and Elephant Shrews 1990
African Primates 1986
Asian Primates 1987
Foxes, Wolves, Jackals and Dogs 1990
Otters 1990
Weasels, Civets, Mongooses and their relatives 1989
Dolphins, Porpoises and Whales 1989
African Elephants and Rhinoceroses 1990
The Asian Elephant 1990
Asian Rhinoceroses 1989
African Antelopes (3 parts) 1988-89
The Kouprey 1988
Rabbits, Hares and Pikas 1990
REPTILES Tortoises and Freshwater Turtles 1989
Crocodiles 1992
INVERTEBRATES Swallowtail Butterflies 1991
Note: See references for full citations. Although part of the SSC network, the Bird Specialist Groups are largely coordinated by the International
Council for Bird Preservation and the International Waterfowl and Wetlands Research Bureau, who are reponsible for a number of bird action plans.
The advantages of a species-based approach lie largely in its
allowing resources to be directed to the most urgent cases,
that is to species known to be most imminently in danger of
extinction. Its disadvantages are that, given existing
knowledge and resources, only a tiny proportion of the
world’s biota can be adequately surveyed to set priorities;
even for those taxa which have been surveyed, adequate
resources are available to implement recovery plans for
only a small proportion of those identified as of high
priority. Anthropocentric prejudice dictates that these are
very largely higher vertebrates (note that no plant and only
three non-mammalian animal SSC Action Plans have been
produced to date). Allocation of extensive resources to the
conservation of a small number of high profile or priority
species may not be the most efficient use of scarce
resources, particularly if conservation measures involve a
large amount of ex situ management, because this does not
benefit any other species.
544
Attempts to reconcile the two approaches centre on the
identification of areas of high diversity and endemism (see
Chapter 15), particularly of threatened species, and the use
of particular species as ‘flagships’ to justify the preservation
of areas of habitat which thereby conserve other species of
lower conservation profile.
The relative merits of these various approaches in
conservation planning, and the degree of intervention
desirable in the management of individual species (in
particular the establishment of captive-breeding groups from
wild populations of critically endangered animal species)
have been, and will continue to be, hotly debated.
This chapter will examine species-based approaches in more
detail, comparing plants and animals and outlining in situ
and ex situ approaches for the two groups.
IN SITU CONSERVATION OF THREATENED PLANT
SPECIES
The most important single way that plant species can ‘be
conserved is by protection of their habitat through control
of land-use. Central to this approach is the network of
protected areas which nearly all countries possess (Chapter
29). However, the overall extent to which these measures
actually preserve wild flora, especially in the tropics where
most species occur, is not known. A survey of 25 (mostly
temperate) countries revealed great variation in the extent
to which listed threatened plant species occurred in
protected areas.
e in New Zealand, of c. 70% of the nationally threatened
species analysed, 71% are in permanent protected areas
managed to benefit the biota, 18% have a low level of in
situ protection, and 11% have no effective protection in
situ (D. Given in litt., 1990).
e In Britain, over 75% of the 317 nationally threatened
plant species are represented in nature reserves (mostly
county wildlife trust reserves) or in Sites of Special
Scientific Interest (SSSI) (L. Farrell in litt., 1990). SSSIs
are in effect ‘quasi protected areas’: they are sites,
usually on private land, designated by the government
conservation service; restrictions may be put on the use
of the land in return for compensation to the landowner;
however, in only a few cases have management
agreements on SSSIs been concluded for nationally
threatened plants.
© In Spain, excluding the Canary Islands, about 35-40% of
the endemic plants grow in protected areas, although
many of these areas are recently designated and have no
effective protection or management (C. Gomez Campo
in litt., 1990).
e In Bulgaria, out of 763 species listed in the National
Plant Red Data Book (Velchev et al., 1984), 38% occur
in protected areas, and 63 (8%) of these are apparently
confined to them.
Current Practices in Conservation
e In Czechoslovakia, all 400 or so species to be included
in the Red Data Book of Higher Plants (Cerovsky et al.,
in prep.) occur in protected areas. This is partly because
the occurrence of threatened species was used as a
priority criterion for creating protected areas. Of the 400
species, at least 118 (30%) have a good measure of
active protection.
e In Poland, out of a list of 339 threatened species of
higher plants, all 308 extant are in protected areas and
an estimated 30% are restricted to them.
e In Australia, almost exactly half of the 3,635 threatened
species are in conservation reserves, with 179 believed
confined to reserves (Table 34.3).
e In Southern Africa, figures are available which relate to
the flora as a whole rather than to threatened species
only. In a recent study (Siegfried, 1989), it was found
that out of the 582 publicly-owned protected areas,
covering 6% of the region, complete plant lists were
available for 52 and partial plant lists for 153. An
analysis of this and other information found 34% of
Southern Africa’s 23,300 known vascular plant species
in reserves. Further analysis, using known plant
distributions, led to a prediction that some 74% of the
Tegion’s vascular plant species were represented in
nature reserves. Of the rich and endangered fynbos
flora, the prediction was of 99% coverage, although the
author says that this is "almost certainly too high"
(Siegfried, 1989). Nevertheless the region’s nature
reserves clearly protect a very high proportion of the
flora and by implication of the threatened species.
These figures indicate that in countries where there have
been long-standing programmes to identify and conserve
threatened plants, some degree of success is possible.
However, in other countries surveyed no threatened plants
are known to be protected in conservation areas.
The information also shows that very often the large
protected areas which receive the most attention, such as
national parks, are not the most important sites for
Table 34.3. Threatened plant species in protected areas in Australia
ADEQUACY OF
RESERVATION
Reserved taxa Endangered Vulnerable
A 4 53
B 38 143
Cc 11 163
Total reserved 53 359
All listed taxa 180 661
% reserved 29.4 54.3
CONSERVATION STATUS
Rare Poorly known Total
218 52 327
45 32 258
625 449 1248
888 533 1833
1173 1537 3635
75.7 34.7 50.4
Source: J.H. Leigh in litt., 18 September 1991, updating Briggs, J.D. and Leigh, J.H. 1988. Rare or Threatened Australian Plants. Australian
National Parks and Wildlife Service.
Notes: ‘Poorly known’: taxa suspected, but not definitely known, to belong to the categories Extinct, Endangered, Vulnerable or Rare; other
categories as used by WCMC and defined on p.234. ‘Reserved taxa’ column: A = 1,000 plants or more are known to occur within a conservation
reserve(s); B = fewer than 1,000 plants are known to occur within conservation reserve(s); C = although recorded from a reserve, the population
size is unknown.
545
3. Conservation and Management of Biodiversity
conserving threatened or endemic plants. Smaller, less well
known sites may be more important. For example in Spain,
excluding the Canary Islands, no more than 5% of the
nationally threatened plants occur in national parks. In the
UK, the small and private nature reserves of the county
wildlife trusts are widely recognised as more important for
protecting threatened plants than the network of National
Nature Reserves.
However, presence in a designated protected area is itself
no guarantee of survival. Levels of protection are often
inadequate or non-existent. Moreover, even in areas under
active protection, appropriate management to maintain
viable populations of threatened plant species may not be
carried out for a variety of reasons - protection of a site
from external disturbance may be enough in some
circumstances, such as large reserves in tropical forests, but
in others, such as temperate grasslands, active management
may be essential.
IN SITU CONSERVATION OF CROPS AND WILD
RELATIVES OF CROPS
As well as conserving threatened species, protected areas
could potentially play a vital role in the conservation of
plant resources which may be of more immediate benefit to
mankind. Two of the most important categories of these are
traditional crops and wild relatives of crops. Traditional
crop material is defined here as pre-20th century varieties
and ‘land races’ which are diverse collections of primitive
types of domesticated material and related weeds.
It is widely accepted that genetic erosion of crop gene pools
continues today at a rapid, albeit unquantifiable rate (Anon.,
1991b). There are a number of causal agents that fuel
genetic erosion and their effects have intensified over the
last decade. These factors include agricultural
mechanisation, spread of uniform hybrids, and habitat
destruction such as forest clearing and urbanisation. The
rates of genetic erosion are greatest in areas with the most
fertile and most easily mechanised agricultural lands,
especially near urban centres and markets. It is in
impoverished and marginal areas, such as mountainous
uplands, where traditional varieties are still grown and are
sufficiently relied upon to allow for their careful
conservation. The crop gene pools subject to active
breeding programmes are among those that have been most
depleted (Fowler and Mooney, 1990).
In terms of conservation requirements, crops and crop
relatives can be divided into two very different groups:
annuals and perennials. Seeds of annuals can be conserved
ex situ through cold storage. However, for many
perennials, ex situ conservation is not workable because of
short-lived or recalcitrant seed, the limitations of meristem
culture and reproduction requirements which are difficult to
meet in field gene-banks and laboratories (Ingram, 1984).
Traditional varieties and wild relatives of crops differ in
their conservation requirements because wild species are
evolving within natural ecosystems while land races and
other primitive material are products of human practices
and modified habitats.
Recent work on the structure of crop populations
(Zimmerer and Douches, 1991) and on wild relatives of
546
crops describes a fine mosaic of genotypic variation. This
genetic diversity is difficult to capture and maintain even in
networks of protected areas and comprehensively developed
gene-bank collections (Brush, 1991). Probably much less
than 10% of the alleles in the gene pools of the major crops
are currently contained in protected areas. The prospects of
maintenance of the genetic variation within these
populations, with pressures for expanded uses of protected
areas and a poor technical and institutional base for
management, are doubtful, especially in view of the fact
that the great majority of crop genetic resources are in the
tropics and in countries with relatively. ineffective
programmes of protected area planning and management.
Great expansion of such programmes is required in order to
avoid accelerating and irreversible loss of potentially
valuable genes.
Levels of effectiveness of in situ conservation of plant
genetic resources
Virtually all protected areas and landscapes have some
populations of some species of economic importance which
are more or less adequately protected, at least in the
short-term. However, in virtually all, substantially increased
monitoring and management programmes are necessary in
order to avert losses of rarer genes and other potential
genetic resources. There are three general levels of in situ
conservation of plant genetic resources which are described
below.
Level I represents largely unplanned coverage through
ecosystem conservation. It cannot be assumed that there will
be adequate coverage within the reserve to maintain viable
populations of plant species with genetic resources over the
long-term.
Level II requires the planning and design of protected areas
with use of distribution data for species with genetic
resources. Many of the ‘genetic reserves’ (Jain, 1975)
involve this level of conservation. Management for
particular species and associated successional phases is
usually necessary. Species that are monitored and managed
under programmes of ecosystem coverage can also have
level II conservation.
Level III involves site-specific monitoring, management and
procurement for particular levels of conservation for
specific "functional population units" (Solbrig, 1991). For
this level of conservation to be attained, population viability
thresholds must be set with prescriptions for maintenance of
intra-specific variation and rarer alleles.
In both natural and well-protected populations, there is a
constant flux of gene frequencies with some alleles
becoming rare or disappearing. In protected areas, the
natural and human-induced dwindling of populations can
cause the narrowing of the base of variation and subsequent
loss of potentially valuable genetic resources. In order to
maintain rarer alleles or possible adaptive complexes
associated with certain environments and selection factors,
further requirements for larger and sometimes additional
populations must be set.
With species for which there is no major concern for
maintenance of potential genetic resources, level I
conservation is often adequate. For in situ conservation of
crop genetic resources, both traditional varieties and wild
species, /evel I] is always necessary to assure capture of
desirable percentages of alleles and /evel J/] is necessary for
long-term security as well as procurement for extended
programmes of plant breeding.
The status of most populations with genetic resources is still
poorly determined and the development of new theory and
techniques for determining conservation requirements are
necessary. There are a number of technical issues which
must be addressed before protected areas can become
effective for the conservation and procurement of genetic
resources. Desired levels of conservation of less common
genes should be determined. Access to and ease of
procurement of the genetic resources of wild species must
be effectively regulated and in some cases expanded.
Regulation and expansion of the systems of distribution of
the germplasm taken from protected populations are
inevitable (Kloppenburg and Kleinman, 1988).
The text below will discuss the current status of the three
levels of in situ conservation of crops and wild relatives of
crops in terms of three categories: traditional land-tenure
and farming systems; genetic reserves and other locally and
nationally managed protected areas; internationally
monitored protected areas.
Traditional and
conservation
in-farm programmes of in situ
Areas with traditional land tenure and farming systems
provide a basis for in situ conservation. These areas may be
designated protected areas or they may be non-protected
areas where key aspects of traditional farming systems are
supported and maintained.
Genetic conservation requires recognition of the
interrelationships of genotypic and allelic diversity and the
functioning of the agroecosystem as a whole. The existing
variation has co-evolved within a mosaic of agroecosystem
zones. It is especially important to know the specific
environmental conditions necessary for conservation of
narrowly adapted land races.
In considering the requirements for in situ conservation of
crops, agricultural communities and systems can be placed
in the following categories:
© highly traditional agricultural mosaic which has only
been moderately altered through rising human population
and new technologies,
traditional agricultural mosaic under stress and with
rapid rates of loss of primitive material and the diversity
of selection factors,
comparatively recent, pioneer communities with limited
local knowledge and with recently introduced cultivated
material.
The first generally holds the most crop genetic resources
while the second has the greatest rates of genetic erosion.
In order to design and maintain effective in-farm
conservation programmes, interrelationships between socio-
547
Current Practices in Conservation
economic systems and the structure of crop populations
must be ascertained. Gender may be an important factor.
Women often play key roles in farm conservation of land
races, especially where they have traditionally been the
selectors of seeds for planting. For example, women in a
number of traditional Ethiopian societies pass knowledge of
seed selection from mother to daughter (Marie Dulude,
pers. comm.) and in a village in Liberia women maintain
112 varieties of rice, matched to particular micro-
environments, slope categories, insolation, and soil types
(Thomasson, 1991).
The traditional agroforestry system practised on Fergusson
Island, Papua New Guinea, serves as an example of in situ
conservation of traditional varieties of crops within a
particularly biodiversity-rich setting (Flavelle, 1990).
Because of low human populations, mountainous terrain,
and distance to market, cash crop ventures have so far been
unsuccessful. The system revolves around the growing of
yams, principally Dioscorea esculenta and D. alata. Yams
are the basis of the subsistence economy but it is their
cultural importance that may ultimately prevent them from
being replaced by introduced food crop species. Yam seeds
are inherited through the matrilineage; they are exchanged
as gifts at funeral feasts and other occasions; they are the
focus of magic ritual and myths. Other tubers which
farmers have grown for generations but which hold less
status than yams include: sweet potato, Ipomea sp.,
Colocasia esculenta, and manihot, Manihot esculenta.
Prominent food trees observed in the system include mango,
Mangifera sp., Citrus sp., Szigium sp., Carica sp., banana,
Musa sp., and Arctocarpus sp., and Ficus sp., Cocos
nucifera, and sago, Metroxylon sp. Every time that a
community opposes intrusion of the cash economy and
environmentally damaging activities, it is a form of in situ
conservation. Such approaches may not be permanent but
can be effective over the medium- and long-term.
There is currently a very poor institutional base for more
coordinated in-farm conservation programmes. The
International Agricultural Research Centres (IARC) manage
information on the major food crops, however, they have
less expertise in integration of information about the
ecosystems, societies and cultures that have created and
now maintain these traditional varieties.
Almost all of the current in-farm crop conservation
programmes have been initiated by small institutes and
NGOs. These programmes tend to emphasise research,
education, technical advice, and credit schemes that support
traditional farming systems and have shown particular
success where they have been able to link conservation
interests with locally-driven rural development.
The Rural Advancement Fund International (RAFI)
promotes the maintenance of land races and low-input
farming and attempts to channel funding to local NGOs
involved in crop conservation. RAFI has developed a
training kit on community plant breeding, and local seed
banking by maintaining living stock. In Ethiopia and
Zimbabwe, the programme has been established through the
umbrella organisation, Seeds for Survival. The Southeast
Asia Regional Institute for Community Education
(SEARICE) is working in the Mekong Delta and the
3. Conservation and Management of Biodiversity
Philippines. CLADES is a South American NGO promoting
sustainable agriculture, including the conservation of land
races and wild crop relatives.
In Ethiopia, the Seeds for Survival programme is working
with the Plant Genetic Resources Centre in Addis Ababa to
reintroduce traditional varieties and land races to farmers.
During droughts, farmers were forced to eat their stocks of
seeds. PGRC responded by making extensive collections
and is now reintroducing land races to farmers, establishing
research plots, monitoring productivity, and establishing
training and support programmes for farmers.
In the developed countries, there is some in situ
conservation in historic farms and other protected cultural
landscapes as well as through the networks of seed
conservationists. Most efforts are organised at the
grassroots level.
In order to implement long-term programmes with the dual
mandate of monitoring genetic resources in agroecosystems
and supporting farmer-driven development, national
institutes and NGOs require greatly increased and more
stable funding bases. For Latin America, Montecinos and
Altieri (1991) stated that, "over 50% of the initiatives
known by the authors do not have their own staff or budget,
but must borrow from other projects, and strongly rely on
help from farmers and other local sources. Among those
that have received financial support, again over 50% have
been working with less than US$5,000 per year. Over 60%
of the programmes have done collection work, but do not
have money to set up simple and adequate storage facilities
or, even worse, to do a systematic monitoring of
multiplication of material and performance tests in the field.
This is, in fact, one very strong reason for diversity loss
and for not seeing many breeding initiatives at the farm
level."
There are major questions about the effectiveness of
virtually all of the in situ conservation programmes for
crops. At the local level, there are overlapping and jumbled
objectives and many farmer organisations are more
concerned with the economic benefits of producing their
own seed than with conservation of genetic diversity.
Genetic reserves and other nationally or locally managed
protected areas
Genetic reserves and managed protected areas often provide
adequate levels of in situ conservation though there are
often insufficient inventory data and management expertise.
The institutional bases for all of the locally managed
programmes of in situ conservation of crop relatives are
remarkably weak and, after nearly a decade of negotiations,
there are still no national or international frameworks
funding in situ conservation of wild relatives of crops.
Genetic reserves (Jain, 1975) have been established for the
procurement of seed and other forms of germplasm for
thousands of years. In recent decades, these reserves have
often been established for single and multiple crop relatives.
The major problem with the current networks of genetic
reserves is that the sites are often too small and do not
include population levels that are adequate for maintenance
548
of fitness and of rarer genes, nor have adequate potential
for a diversity of management treatments.
In less densely populated regions with remaining forest,
extractive reserves as in Amazonia could be managed for
the genetic resources of wild relatives of crops. These
reserves have been established for the benefit of local
people to gather Brazil nuts, Bertholletia excelsa, and other
species with commodity value. Some wild or semi-
domesticated species thrive in more altered habitats. Fruit
crops such as papaya, Carica papaya, bacuri, Platonia
insignis, guava, Psidium guajava, and ciruela, Bunchosia
glandulosa, have wild populations in more disturbed and
open forest mosaics dominated by second-growth forest.
Cupuaca, Theobroma grandiflorum, is a wild relative of
cacao that is planted from seed in backyards in Amazonia,
particularly in Para (Smith and Schultes, 1990).
In other types of protected area, advances in management
of genetic resources have been limited. While protected area
managers worldwide have become increasingly aware of
issues of genetic resources and maintenance of genetic
diversity, these are generally afforded lower priority than
other management concerns. Moreover, traditional farming
and agroforestry have tended to be suppressed within many
protected areas, with a concomitant increase in rates of
genetic erosion of crop plants.
There is still inadequate information on the status of crop
relatives in protected areas. The level of funding for this
type of highly technical and relatively expensive
inventorying, monitoring and management may actually
have declined, in real terms, in most of the world over the
last decade. Even in developed countries attention is
directed to threatened habitats and threatened species and
few resources are available for monitoring populations of
crop relatives, although some work has been done (e.g. on
wild cranberries Viburnum spp. in protected areas in the
mid-Atlantic States of the USA). The most effective
technical linkages between programmes in the developing
countries have been forged by FAO (Palmberg and
Esquinas-Alcazar, 1990). However, such national
programmes have tended to focus on timber species rather
than food crops.
Biosphere reserves, World Heritage Sites and other
internationally monitored protected areas
The internationally monitored protected areas, which are
principally biosphere reserves and World Heritage Sites,
hold the greatest promise for adequate inventorying,
monitoring, managementand procurementthough respective
levels of national support are too often as weak as with
other protected areas.
Biosphere reserves will play an increasingly key role in the
conservation and utilisation of wild relatives of crops
because of institutional potentials for monitoring and
international exchange of information (Ingram and
Williams, 1984; Ingram, 1990b). The concept has not been
effectively utilised for traditional crops. The network is
very new (Batisse, 1982; Vernhes, 1989) and most reserves
are still poorly inventoried.
There has been no systematic documentation of the crop
varieties, land races and other primitive material in either
the transition areas or in the buffer zones of biosphere
reserves or within World Heritage Sites. There has been
only cursory reporting of agricultural activities within and
on the edge of biosphere reserves and World Heritage Sites,
both natural and cultural, have rarely been inventoried for
crop varieties.
The most impressive example of in situ conservation of
crops in an internationally monitored protected area is that
of Niger’s Air and Ténéré National Nature Reserve which
maintains a range of traditional and contemporary garden
types and primitive, traditional and more recent crop
material for the gene pools of sorghum, Sorghum spp.,
pearl millet, Pennisetum spp., barley, Hordeum sp., and
wheat, Triticum sp. as well as for agroforestry: Acacia
spp., Olea sp. and Ziziphus spp. (Ingram, 1990a). The area
is at an ecological crossroad in terms of material and
farming practices from North Africa and the Sahel. Despite
the present maintenance of these gardens within the
framework of protected area management, progressive
desertification due largely to climatic change, could destroy
prospects for long-term in situ conservation.
Only a very small portion of the 300 biosphere reserves
(see Chapter 29) which are part of UNESCO’s Man and the
Biosphere Programme are in centres of crop origin or in the
regions of high species diversity, such as the humid tropics.
Based on cursory documentation (MAB, UNESCO files,
Paris), well over 20% of the biosphere reserves have
populations of the relatives of the major crops (Table 34.4);
with better documentation and complete inventories of plant
species in current biosphere reserves, this percentage is
expected to exceed 50%.
There are, however, virtually no distribution data available
and consequently no assessment of the status of populations
in terms of core, buffer and transitional zones. The
information on wild relatives in World Heritage Sites is
even more cursory. Consequently, there is very little level
III in situ conservation anywhere though these areas have
the best institutional potentials.
The actual status of wild relatives in many biosphere
reserves will remain contentious, especially with changing
political and administrative contexts. For example, the
status of the many crop genetic resources, such as Malus —
spp. and Prunus spp., which provided the original focus for
a number of reserves in Soviet Asia, has not been reviewed
in recent years. The Sierra de Manantlan Biosphere Reserve
in Mexico was recently established for the conservation of
the genetic resources of wild corn, Zea spp., though the
requirements for Jevel III in situ conservation have still
barely been met.
Conclusions
Only a tiny portion, well under 10%, of the total species
and allelic diversity of the major crop gene pools are
currently maintained in situ and of this only a small portion
is conserved at levels adequate to withstand threats over the
long-term as well as needs for germplasm for breeding
programmes. Genetic erosion continues and in many cases
549
Current Practices in Conservation
is accelerating both outside and within protected areas.
However, there are inadequate inventory and monitoring
data to determine rates. Greatly increased programmes of
protected area planning, monitoring, and management are
needed within the next five years in order to cause a
significant reversal of the accelerating rates of genetic
impoverishment.
INSTITUTIONS INVOLVED
CONSERVATION OF PLANTS
IN EX SITU
Botanic Gardens
The single most important type of institution involved in ex
situ conservation of wild plants is the botanic garden. There
are over 1,500 botanic gardens worldwide, of which about
800 are believed to be currently active in plant
conservation, although this number is steadily increasing.
Botanic gardens worldwide contain well over three million
accessions between them. This total illustrates the
remarkable capacity of the network of botanic gardens for
the cultivation of plants and their potential contribution to
plant conservation. However, many botanic gardens are
poorly financed and badly organised, or only weakly
supported within their institutions or by their governments,
and a large percentage of plants grown is of low
conservation priority.
There is a considerable imbalance in the global distribution
of botanic gardens. Europe has 532 botanic gardens, but
Africa has only 82 and South America 66. Most tropical
countries, where most of the world’s flora resides, have few
botanic gardens (Table 34.5) and most of these are poorly
developed or funded, although most of the new botanic
gardens that have been created or planned in recent years
are in tropical countries, which have large floras. Indeed,
many gardens in temperate countries have ceased to play
any significant role in conservation or research and retain
only educational significance, whereas most new tropical
gardens have been created primarily as centres for plant
conservation. For example, the Conservatoire et Jardin
Botanique de Mascarin, Réunion Island, founded in 1987,
contains over 60% of the island’s rare and endangered
flora.
The collections maintained in botanic gardens are very
diverse. Particular groups, such as orchids, succulents,
bromeliads, bulbous species and temperate trees, are
particularly well represented in cultivation, as Table 34.6
shows. Collections of tropical woody species are, however,
less well represented. In general, the floras of tropical and
sub-tropical continental countries are less commonly grown
than those of temperate countries and oceanic islands.
Efforts to coordinate the activities of botanic gardens at an
international level are undertaken by Botanic Gardens
Conservation International (BGCI). The purpose of the
Secretariat is to disseminate information to promote and
coordinate the ex situ conservation of threatened wild
plants. It also provides technical guidance, data and support
for botanic gardens in almost 100 countries and assists and
promotes the development of botanic gardens and their plant
conservation programmes. BGCI has a worldwide
membership of 317 botanic gardens (Table 34.5).
3. Conservation and Management of Biodiversity
Table 34.4 Biosphere Reserves with higher levels of documentation of wild
Algeria
Argentina
Australia
Austria
Brazil
Bulgaria
Cameroon
Canada
Central African
Republic
Chile
China
Colombia
Costa Rica
Czechoslovakia
Denmark
Egypt
Ecuador
France
Germany
relatives of crops and forage species
NAME OF
PROTECTED AREA
Parc national de Tassili
N’Ajjer
El Kala PN
Reserva Ecolégica de
Nacufian
Croajingolong
Lobau Reserve
Benin
Reserva de la biosphere de
la Pendjari
Vale do Ribeira and Serra
DA Graciosa Biosphere
Reserve
Reserve Boatine
Reserve Ouzounboudjak
Parc national de Waza
Parc national de la Bénoué
Waterton Lakes National
Park
Bamingui-Bangoran
Conservation Area
Parque Nacional Juan
Fernandez
Reserva de la Biosfera La
Compana-Pefuelas
Changbai Mountain
Biosphere Reserve
Dinghu Nature Reserve
Fanjings han Mountain
Nature Reserve
Shennongjia Biosphere
Reserve
Cinturon Andino Cluster
Biosphere Reserve
Sierra Nevada de Santa
Marta
Reserva de la Biosfera de
la Amistad
Cordillera Volcanica
Central
Palava Protected
Landscape Area
Northeast Greenland
National Park
Omayed Experimental
Research Area
Galapagos - Archipiélago
de Colon
Réserve de la biosphére de
PN des Cévennes
Steckby-Loedderitz Forest
Nature Reserves
Vessertal Nature Reserve
Middle Elbe Biosphere
Reserve
Southeast Riigen
Biosphere Reserve
BIOGEOGRAPHIC
PROVINCE
Sahara
Mediterranean sclerophyll
Monte
Eastern sclerophyll
Central European
highlands
West African woodland
and savanna
Serra Do Mar
Balkan Highlands
Balkan Highlands
West Africn woodland and
savanna
West African woodland
and savanna
Rocky Mountains
West African woodland
and savanna
Southeastern Polynesian
Chilean Sclerophyll and
Chilean Nothofagus
Manchu-Japanese Mixed
Forest
South Chinese Rainforest
Chinese Subtropical Forest
Chinese Subtropical Forest
Northern Andean
Venezuelan Dry Forest
Central American
Central American
Middle European Forest
Arctic desert and icecap
Sahara
Galapagos Islands
Atlantic
Middle European Forest
Middle European Forest
Middle European Forest
Middle European Forest
550
AREA OF
RESERVE (ha)
7,200,000
76,438
11,900
101,000
1,000
880,000
1,615,000
1,226
2,575
170,000
180,000
52,597
1,622,000
9,290
17,095
217,235
1200
41,533
29,400
855,000
731,250
584,592
144,393
8,017
70,000,000
1,000
766,514
323,000
3,500
17,500
18,640
BETTER DOCUMENTED GENERA
WITH GENETIC RESOURCES
Olea spp.
Olea sp., Pistacia sp.
Prosopis spp.
Acacia spp.
Pyrus sp.
Acacia spp.
Prunus sp.
Prunus sp.
Vaccinium sp.
Oryza sp., Sorghum sp.
Pennisetum spp., Prosopis spp.
Allium sp.
Prosopis sp.
Rubus spp.
Ribes sp.
Panax spp.
Castanopsis sp.
Vaccinium spp.
Prunus sp., Brassica sp., Ribes
sp., Malus sp., Prunus sp.,
Rubus sp., Vitis spp., Panax sp.,
Allium spp., Avena spp.,
Sorghum sp. land races
Juglans sp.
Acacia spp.
probably Persea spp.,
Lycopersicon spp.
probably Persea spp. and
Lycopersicon spp.
Prunus sp., Avena sp., Solanum
sp.
Vaccinium sp.
Gymnocarpus sp.
Lycopersicon sp.
Castanea sp., Vaccinium sp.
Pyru sp.
Vaccinium spp.
Malus sp., Pyrus sp.
Allium sp., Malus sp., Fragaria
sp., Vaccinium spp.
Current Practices in Conservation
Table 34.4 Biosphere Reserves with higher levels of documentation of wild
relatives of crops and forage species (continued)
Greece
Guatemala
Indonesia
lran
Italy
Kenya
Mali
Mauritius
Mexico
Mongolia
North Korea
Pakistan
Peru
Philippines
Poland
Romania
Senegal
Spain
Gorge of Samaria National
Park
Mount Olympus National
Park
Tikal National Park
Cibodas Biosphere Reserve
Komodo Proposed National
Park
Tanjung Puting Proposed
National Park
Gunung Leuser Proposed
National Park
Siberut Nature Reserve
Arasbaran Protected Area
Gano Protected Area
Hara Protected Area
Miankaleh Protected Area
Touran Protected Area
Foret Domaniale du Circeo
Mount Kulal Biosphere
Reserve
Amboseli Biosphere
Reserve
Parc national de la Boucle
du Baoucle -
Macc habée-Bell Ombre
Nature Reserve
Montes Azules
Reserva de la Biosfera de
Sian Ka‘an
Reserva de la Biosfera
Sierra de Manantlan
Great Gobi Biosphere
Reserve
Mount Paekdu Nature
Reserve and Biosphere
Reserve
Lal Suhanra National Park
Reserva del Noroeste
Palawan Island Biosphere
Reserve
Babia Gora National Park
Pietrosul Mare Nature
Reserve
Retezat National Park
Rosca-Letea Reserve
Forét classée de Samba
Dia
Parc national du
Niokolo-Koba
Reserva de Grazalema
Reserva de la Biosfera de
Donana
Reserva de la Biosfera del
Urdaibai
Parque Natural Del
Montseny
Mediterranean Sclerophyll
Mediterranean Sclerophyll
Campechean
Java
Lesser Sunda Islands
Borneo
Sumatra
Sumatran
Caucaso-lranian Highlands
Anatolian-lranian Desert
Anatolian-lranian Desert
Caucaso-lranian Highlands
Iranian Desert
Mediterranean Sclerophyll
Somalian/Lake Rudolf
Somalian Grasslands
West African woodland
and savanna
Mascarene Islands
Campechean
Campechean and
Yucatecan
Madrean-Cordilleran
Gobi Desert
Manchu-Japanese Mixed
Forest
Thar Desert
Equadorian Dry Forest
Philippines
Middle European Forest
Middle European Forest
Middle European Forest
Pontian Steppe
West African woodland
and savanna
West African Woodland
and savanna
Mediterranean Sclerophyll
Mediterranean Sclerophyll
Iberian Highlands
Mediterranean Sclerophyll
Source: table compiled by G.B. Ingram, based on MAB/UNESCO files.
Notes: ' This biosphere reserve was the first set up primarily for the protection of wild corn, teosinte, Zea diploperennis.
In recent years new national botanic garden organisations
have been formed in many countries, such as Brazil and
Australia, with regional groupings of the International
551
4,840
4,000
57,600
140,000
70,000
52,000
49,000
85,686
68,800
1,000,000
3,260
700,000
283,200
771,000
3,611
331,200
528,000
139,577
5,303,172
60,000
31,355
226,300
1,150,800
1,741
3,068
20,000
18,145
756
913,000
32,210
77,260
22,500
17,372
Olea sp.
Allium sp.
Acacia sp., Lycopersicon sp.,
Persea sp.
Vaccinium spp.
Jpomoea spp.
Durio sp.
Musa spp., Citrus spp.
Durio spp., Musa spp.
Juglans sp.
Olea sp., Prunus sp., Pistacia sp.,
Acacia spp.
Acacia sp., Prosopis sp.
Punica sp., Rubus sp.
Hordeum sp.
Pistacia sp.
Olea sp., Acacia sp.
Acacia spp.
Acacia spp.
Coffea sp.
Persea sp.
probably Persea sp.
1
Hordeum sp., Allium sp.
Ribes spp., Rubus spp., Prunus
sp., Vaccinium spp., Viburnum
sp., Allium sp.
Acacia spp.
Acacia spp.
Durio spp.
Allium sp.
Allium spp.
Juglans spp.
Vitis spp.
Acacia spp.
Acacia spp.
Ceratonia sp.
Olea sp.
Castanea sp.
Prunus sp.
Association of Botanic Gardens (IABG) existing in, for
example, Europe, the Mediterranean region, Latin America
and Asia.
3. Conservation and Management of Biodiversity
Table 34.5 Number of botanic gardens and known cultivated accessions in
botanic garden collections
COUNTRIES NO. OF NO. OF NO. OF COUNTRIES NO. OF NO. OF NO. OF
BOTANIC BGCI ACCESSIONS BOTANIC BGCI ACCESSIONS
GARDENS MEMBERS _IN CULTIVATION GARDENS MEMBERS _IN CULTIVATION
ASIA NORTH AND CENTRAL AMERICA (continued)
Bangladesh 2 0) 700
China 66 4 56,278 Canada 18 7 67,374
Hong Kong 4 3 1,200 Cayman Is 1 1 =
India 68 7 86,259 Costa Rica 2 2 4,000
Indonesia 5 4 69,840 Gabe 8 3 7,550
ee z £ eae Dominica 1 1 750
Iraq 1 oO 230 Dominican Rep. 1 fe) -
Israel 7 3 ee El Salvador 1 1 3,500
Japan 59 (0) 72,560 Grenada 1 0 =
Korea, DPR 1 0 3,140 Guadeloupe 2 1 100
Korea, Rep 5 0 10,000 Guatemala 1 1 700
Malaysia 9 5 3,872 Honduras 2 1 764
Mongolia 1 v Z Jamaica 4 oO 1,557
Myanmar 2 O : Martinique 3 0 200
Nepal 1 u etl Mexico 30 7 8,650
Pakistan 5 10) 430 Nicaragua 1 0 ys
Philippines 9 2 16,829 Panama 1 fo) S
Saudi) Arabia 2 ° : Puerto Rico 4 0 3,150
Singapore 1 u 3,000 St Vincent 1 1 é
Sri Lanka 6 6 7,125 Stinonc 1 0 *
Taiwan 2 1 2,513 Trinidad and Tobago 1 0 -
Thailand 5 10) 2,400 USA 247 47 424,888
MERE) i u Bue (Hawaii) 19 6 25,632
Viet Nam 3 1 Virgin Is (British) 1 1 5,000
USSR {former} Virgin Is (US) 1 ty) 3,000
USSR 160 1 344,744 SOUTH AMERICA
EUROPE Argentina 9 1 18,687
Bolivia 3 1 -
Albania 1 (0) 2,000 Brazil 11 4 20,820
Austria 11 10) 40,300 Chile 9 1 1,967
Belgium 15 3 45,783 Colombia i) 4 3,000
Bulgaria 9 (0) 3,000 Ecuador ei 1 500
Czechoslovakia 34 1 53,817 French Guiana 2 (0) -
Denmark 7 2 48,950 Guyana 2 1 300
Finland 8 3 22,900 Paraguay 1 10) -
France 66 18 171,725 Peru 5 1 -
Germany 73 12 383,470 Uruguay 1 10) 500
Greece 4 2 3,550 Venezuela i 2 1,003
Hungary 17 10) 7,350
Iceland 2 (0) 6,500 eee
olen 8 5 36,500 Australia 60 22 99,752
Italy 48 10 118,432 Fiji 1 o 3
Malta 1 O 8,000 New Zealand : 17 6 28,231
Monaco 1 1 7,000 Papua New Guinea 4 2 6,700
Netherlands 39 5 95,180 SSIEINED |e 4 U 5
Norway 6 3 14,400 Western Samoa 1 oO -
Poland 25 1 54,066 AFRICA :
Portugal 6 1 13,204 Algeria 3 fe} 8,000
(Azores) 3 2 100 Angola 1 fo) 500
(Macau) 1 (0) = Benin 1 fo) =
eects 2 p - Burundi 1 ) -
Romania 10 (0) 42,400 Cameroon 2 1 ;
Spain 8 5 15,900 Cape Verde 1 oO E
{Balearicils) 1 0 ; Céte d'Ivoire 1 0 1,200
(Canary Is) 3 1 10,000 Egypt 5 1 7,550
Sweden 9 3 38,190 Ethiopia 1 (0) 100
Switzerland 22 9 82,020 Gabon 1 1 Z
UK 60 31 217,341 Ghana 3 1 1,000
(Gibraltar) 1 (0) = Kenya 5 1 710
Yugoslavia 32 19) 29,508 Libya 1 (0) 400
NORTH AND CENTRAL AMERICA Madagascar 1 1 5,000
Barbados 2 te) - Malawi 3 2 ile
eliza 1 O - Mauritius 2 2 880
Ranntda 1 0 iu Morocco 2 Oo 1,200
Mozambique 2 10) 2,200
552
Current Practices in Conservation
Table 34.5 Number of botanic gardens and known cultivated accessions in
botanic garden collections (continued)
COUNTRIES NO. OF NO. OF NO. OF
BOTANIC BGCI ACCESSIONS
GARDENS MEMBERS IN CULTIVATION
AFRICA (continued)
Namibia
Nigeria
Réunion
Rwanda
Senegal
Seychelles
Sierra Leone
South Africa
No-+060 25h 02
-OC0O00CONR-
w
fo}
to}
27,582
a
—=
COUNTRIES NO. OF NO. OF NO. OF
BOTANIC BGCl ACCESSIONS
GARDENS MEMBERS IN CULTIVATION
AFRICA (continued)
St Helena 1 (0) -
Sudan 1 ie) 150
Tanzania 2 1 -
Togo 1 1 200
Tunisia 1 {e) -
Uganda 7 {e) 3,320
Zaire 2 2 2,560
Zimbabwe 4 2 3,250
TOTAL 1555 317 3,077,643
Source: International Directory of Botanic Gardens 1990 and BGCI unpublished information.
Most botanic gardens now recognise that priority should be
given to growing plant material of known wild origin and
gradually many gardens are replacing or supplementing
their collections with accessions from known wild sources.
A shift in emphasis from botanic gardens growing wide and
diverse collections of exotic species to the cultivation of the
native flora of their region has gathered momentum.
Figures taken from the BGCI database on the occurrence of
the rare and threatened plants in botanic gardens are given
in Table 34.6. At present the database includes some
29,000 records of rare and threatened plants of around
10,000 taxa in 400 different institutions. This table indicates
that some geographical regions and taxonomic groups are
well represented in cultivation, such as those from China
(63% of the rare and threatened plants are known to be
cultivated) and Macronesia (77% cultivated) and the family
Cactaceae (85% cultivated). Other regions and groups such
as orchids have been less well surveyed for the database
and appear less well represented in cultivation than they
undoubtedly are; Table 34.8 gives some indication of the
large number or orchid species in cultivation. The low
Table 34.6
garden collections
REGION OR NO. OF RARE AND
PLANT GROUP THREATENED TAXA
IUCN SURVEYED
Macaronesia' 557
China 338
New Zealand 230
South Africa 1,051
Australia 1,867
Mascarene? 377
Europe 1,723
USA 3,324
India 927
Cuba 874
Cacti 451
Cycads 105
Conifers 264
Palms 665
Ferns 600
Orchids 986
overall figures for Europe and the USA are disappointing
and suggest that many gardens in these regions have not yet
sufficiently adapted their activities to be able to contribute
significantly to native plant conservation.
The botanic gardens of China show the benefit of a national
strategy for conservation of flora. Their priority is to bring
into cultivation the protected species of China. Botanic
gardens in each province have particular responsibility for
the endangered species of that province (Table 34.7). Other
gardens have specialist collections such as the Institute of
Medicinal Plant Development, Chinese Academy of
Medicinal Sciences, Beijing, which has collected 62 rare
and threatened species of medicinal plants, and the South
China Institute of Botany, Guangzhou, which has 99 of the
130 species of Chinese Magnoliaceae in cultivation (19 of
which are protected), representing well over a third of the
Magnoliaceae worldwide.
In many cases the genetic diversity maintained in the
gardens’ ex situ holdings is inadequate for conservation
Examples of rare and threatened taxa known in cultivation in botanic
NO. KNOWN IN %
CULTIVATION
BOTANIC GARDENS
419 75
211 63
129 56
514 49
893 48
160 42
558 32
890 26
105 13
55 4.5
385 85
81 77
179 68
298 45
73 12
306 31
Source: BGCI database (1991), based on WCMC threatened plants database list.
Notes: ' Canary Is, Madeira, Azores, Salvage Is, Cape Verde are part of Macaronesia but were not included in the survey.
? Mauritius, Rodrigues, Réunion in the Indian Ocean.
3. Conservation and Management of Biodiversity
Rare and endangered plants in cultivation in botanic gardens and
arboreta in China arranged according to province
Table 34.7
PROVINCE NO. OF PROTECTED
SPECIES IN PROVINCE
Jiangxi 52
Zhejiang 55)
Heilongjiang 16
Jiangsu 76
Hubei 55
Hunan 70
Guangdong 12
Guangxi 115
Shaanxi garden 1 37
Shaanxi garden 2 37
Gansu 10
Medicinal plants, Beijing 62
Nanjing 17
NO. OF PROTECTED SPECIES
IN PROVINCE IN CULT.
TOTAL NO. NATIONALLY
PROTECTED IN CULT.
47 47
12 12
38 59
65 65
67 67
29 29
27 27
Source: Shan-An, He, Heywood, V.H. and Ashton, P.S. 1990. Proceedings of the International Symposium on Botanical Gardens. ISBG, 25-28
September 1988, Nanjing. Jiangsu Science and Technology Publishing House, Nanjing, China.
Note: 389 nationally protected species (National Environmental Protection Bureau of China and the Institute of Botany, 1987).
Table 34.8 Some important living collections of orchid species
COUNTRY COLLECTION NO. OF SPECIES
Australia Canberra National Botanic Garden 800
Brazil Sao Paulo Botanic Gardens 1500
Costa Rica Lankester Botanic Gardens 2,000
Cuba Orquideario Soroa 700
France Jardin des Plantes, Paris 500
Germany Heidelberg University Botanic Garden 2,000
Palmengarten, Frankfurt 1,000
India Orchid Research and Development, Arunachal Pradesh 400
National Orchidarium and Experimental Garden, Yercaud =
Gurukula Botanical Sanctuary, Kerala -
Indonesia Bogor Botanic Garden 883
Cibodas Botanic Garden 230
Purwodadi Botanic Garden 546
Bali Botanic Garden 459
Japan Hiroshima Botanical Garden 3,000
Malaysia Serdang (MARDI) 250
(Sabah) Tenom Orchid Centre 453
(Sarawak) Orchid Centre, Kuching =
Mexico Asociacién Mexicana de Orquideologia 550
Papua New Guinea Lipizauga Botanical Sanctuary, Goroka -
UK Royal Botanic Gardens, Kew 4,000
Royal Botanic Garden, Edinburgh 1,500
Glasgow Botanic Garden 1,000
USA New York Botanical Garden 1,000
Smithsonian National Orchid Collection, Washington 2,500
Wheeler Orchid Collection, Bell State University 3,000
The Marie Selby Botanical Garden, Florida 2,000
Source: Various, including J. Stewart, pers. comm. 1990. Royal Botanic Gardens, Kew, UK.
purposes, as the holdings do not constitute representative
samples of the genetic variation of the species.
Frequently a potentially important species is represented by
no more than an accession of only a few specimens.
Furthermore, many current horticultural and management
practices contribute to continuing genetic erosion even of
these small samples. However, there has been recent
recognition in the botanic garden community of the need for
careful genetic management of their accessions to maximise
genetic diversity. This has led a greater number of botanic
554
gardens to define new procedures for maintaining their
collections. Organisations such as BGCI and the Center for
Plant Conservation (St Louis, USA) are publishing
guidelines for ex situ collection management.
International Agricultural Research Centres
The International Agricultural Research Centres (IARC),
supported by the Consultative Group on International
Agricultural Research (CGIAR), have been active in the
international coordination of activities concerned with plant
Table 34.9 Germplasm holdings of |ARCs
IARC
CIAT
Centro Internacional de
Agricultura
Cali, Columbia
CIMMYT
Centro Internacional de
Mejoramiento de maiz y Trigo
Londres, Mexico
cIP
Centro Internacional de La Papa
Lima, Peru
IBPGR
International Board for Plant
Genetic Resources
Rome, Italy
ICARDA
International Centre for
Agricultural Research in Dry
Areas
Aleppo, Syria
ICRISAT
International Crops Research
Institute for the Semi-Arid
Tropics
Hyderabad, India
ITA
International Institute of Tropical
Agriculture
Ibaden, Nigeria
ILCA
International Livestock Centre
for Africa
Addis Ababa, Ethiopia
MANDATE NO. OF
ACCESSIONS
Phaseolus bean, cassava, rice, 66,000
tropical pastures
wheat 70,000
maize
triticale
potato 12,000
sweet potatao
to further the study, collecting, (189,000)!
conservation, documentation,
evaluation, and use of the genetic
diversity of useful plants for the
benefit of people throughout
the world.
barley, 87,000
lentil,
faba bean,
durum wheat,
bread wheat,
kabuli chickpea
sorghum, 96,000
millet,
chickpea,
pigeonpea,
groundnut
cassava, 36,000
maize,
plantain,
cowpea,
soybean,
rice,
yam,
Livestock production systems in 9,000
sub-Saharan Africa
555
Current Practices in Conservation
GERMPLASM HOLDINGS
Phaseolus vulgaris 35,950
other Phaseous spp. 5,111
Manihot esculenta 4,600
*4,000
Manihot (wild spp.) 32
forage legumes 17,982
forage grasses 2,514
maize 10,500
wheat 60,000
potato 5,000
potato (wild spp.) 1,500
sweet potato 5,200
cereals 49,749
food legumes 16,890
forages 19,952
sorghum 31,030
pearl millet 19,796
chickpea 15,564
pigeonpea 11,040
groundnut 12,160
finger millet 2,848
foxtail millet 1,404
proso millet 831
little millet 401
barnyard millet 582
kodo millet 544
sweet potato *1,000
plantain *250
cassava "2,000
yams *1,000
Musa spp *200
cowpeas 15,100
rice 12,000
Bambara groundnut 2,000
soybean 1,500
wild Vigna 810
grasses 1,524
legumes 6,443
browse species 1,429
3. Conservation and Management of Biodiversity
Table 34.9 Germplasm holdings of IARCs (continued)
IARC MANDATE NO. OF GERMPLASM HOLDINGS
ACCESSIONS
IRRI rice 83,000 Oryza sativa 78,420
International Rice Research (Asian rice)
Institute 0. glaberrima 2,408
Manila, Phillipines (African rice)
wild species and species 2,214
hydrids
genetic testers and 208
mutants
taxa in genera related to 21
oryza
WARDA rice 6,000 rice 5,600
West African Rice Development
Asociation
Céte d'Ivoire)
Notes: * (in vitro). The ‘germplasm holdings’ column gives an approximate taxonomic breakdown of the ‘number of accessions’ column; additative
differences will be because of rounding of figures and different data sources. Total holdings worldwide are estimated at 2.6 million, or, allowing
for duplication, 1.3 million unique samples. ' Number of samples collected by IBPGR or with IBPGR support 1974-1989. Three additional IARCs
have no germplasm collections: IFPRI (International Food Policy Research Institute) Washington, DC, USA; ILRAD (International Laboratory for
Research on Animal Diseases) Nairobi, Kenya; ISNAR (International Service for National Agriculture Research) The Hague, Netherlands.
Source: Germplasm data from Anon. nd. Partners in conservation: plant genetic resources and the CGIAR system. CGIAR/IBPGR. IBPGR sample
estimate and accession numbers from van Sloten, D.H. 1990. IBPGR and the challenges of the 1990s: a personal point of view. Diversity 6(2):36-39.
Table 34.10 Status of crop germplasm collections
TOTAL OF ACCESSIONS TOTAL OF ACCESSIONS
CULTIVATED WILD CULTIVATED WILD
MAJOR CEREALS FOOD LEGUMES
Triticum spp. 567,190 11,986 Glycine max 129,043 10,342
Aegilops spp. -- 14,937 Arachis hypogaea 62,981 4,769
Oryza sativa 242,599 90,814 Phaseolus vulgaris 96,341 16,994
pos MVE 208,227 53 Phaseolus lunatus 12,197 1,168
eae fee pee eee Boca Phaseolus coccineus 3,543 807
“ 4 ‘ Vigna unguiculata 31,180 52
MILLETS Vigna radiata 19,103 20
Pennisetum glaucum 28,609 3,366 Vigna subterranea 2,110 a
Setaria italica 17,234 -- Cicer arietinum 49,176 150
Eleusine coracana 12,363 352 Cajanus cajan 16,463 176
Paspalum scrobiculatum 1,515 -- Vicia faba 20,739 117
ROOTS AND TUBERS Lens culinaris 20,252 54
om pean Bean Lupinus mutabilis 6,267 --
‘olanum spp. F , - **19 368
Jpomoea batatas 20,160 1,853 mai se Pe wie 1g
Dioscorea spp. 10,493 78 eee :
Manihot esculenta 24,219 = Psophocarpus tetragonolobus 5,725 408
Source: International Board for Plant Genetic Resources, 1991.
Note: Information as made available to IBPGR conservation database. * Refers to Teosinte spp. and Tripsacum spp. ** Refers to Lupinus spp.
other than L. mutabilis, L. albus and L. luteus.
resources, particularly gene banks. The CGIAR was specific crops and actively collect on a worldwide basis (see
founded in 1971 and consists of a consortium of donor Table 34.9). The collection efforts of the CGIAR network
countries, foundations and development banks, sponsored were initially focused on crop plants and were based on the
by the World Bank, UNDP and FAO. The establishment of economic importance of the crop, the quality of existing
this international network was motivated by international collections and the degree of threat to the crop.
concern over the problems of genetic erosion in cultivated
species and the loss of related wild species of flora. Perhaps the most important of these IARCs is the
International Board for Plant Genetic Resources (IBPGR) in
At present there are 13 [ARCs supported by the CGIAR. Rome, Italy. Established in 1974, the IBPGR does not store
Most of these centres have specific responsibilities in crop any germplasm itself but has a coordinating role in setting
varietal development and germplasm conservation. A few priorities and creating a network of national programmes
of these centres also serve as an international base for and regional centres for the conservation of plant
556
germplasm. It has provided training facilities, supported
research into techniques of plant germplasm conservation,
sponsored numerous collection missions and provided small
amounts of financial assistance for conservation facilities in
the developing world.
IBPGR has achieved many of its original objectives with
regard to collection of germplasm of many of the major
crops of the world. With IBPGR assistance the 13 IARCs
and 227 seed banks in 99 countries now hold 90% or more
of the known land races of such crops as wheat, corn, oats
and potatoes. The IARCs have an estimated 465,000
accessions in storage, amounting to 35% of unduplicated
world holdings (Van Sloten, 1990) (See Table 34.9). Data
on accessions of cultivated and wild crops, made available
to the IBPGR conservation database by national and other
centres, are presented in Table 34.10. These figures, which
do not claim to be comprehensive, in general show that
collections have grown significantly (by 190% and 20% in
the case of Zea mays and Oryza sativa) over the past
decade.
The network has also successfully encouraged many
national programmes and assisted in many scientific and
educational programmes so that now IBPGR has links with
over 500 institutes in some 106 countries.
The CGAIR network has recently been subject to
controversy. Critics maintain that the organisation is guided
too firmly by the industrial interests of the developed world
(this controversty is not discussed here, but is well
reviewed by Kloppenburg, 1988). FAO has recently
renewed its efforts in this area of conservation, due in large
part to the controversy surrounding the activities of the
IBPGR and the international network it coordinates. The
result of this renewed effort by FAO was the formation of
a new commission called the Commission on Plant Genetic
Resource (CPGR) and the drafting of a legal instrument
known as the International Undertaking on Plant Genetic
Resources (the Undertaking).
The Undertaking called for an international germplasm
network to be established under the auspices of FAO. It
lays out the duty of each nation to make all plant genetic
material freely available and calls for the development of a
procedure under which a germplasm conservation centre
could be established by the FAO. It further provides that
the IBPGR was to continue in its role of coordination but
that it would do so under the supervision of FAO.
The CPRG meets every two years to review progress in
germplasm conservation. The commission held its first
meeting in Rome in 1985, where much of the discussion
focused on concerns with the Undertaking and in situ
preservation, which to a large extent had been ignored by
the IBPGR.
The controversy which the CPGR and the Undertaking were
established to resolve has bedevilled the meetings of the
CPGR. As a result this initiative has been able to achieve
very little so far. Consequently the IBPGR has remained
fairly autonomous and continues to be the main body
coordinating at an international level conservation efforts in
this area.
557
Current Practices in Conservation
TECHNIQUES FOR EX SITU PLANT
CONSERVATION
Maintenance of ex situ populations of plants carried out by
a variety of institutions, including botanical gardens,
forestry institutes and agricultural research centres, involves
three important techniques which will be outlined here.
These are field gene banks, seed banks, and in vitro storage
methods.
Field Gene Banks
A field gene bank is an area of land in which collections of
growing plants have been assembled including as many
individuals of one species as possible in order to maintain
the widest practicable range of genetic diversity. This
ensures that plant material is conserved and available for
breeding, reintroduction, research and other purposes. Field
banks are particularly appropriate for long-lived perennial
trees and shrubs which cannot be adequately conserved in
the wild and which may take decades to produce seeds; they
thus have particular importance in forestry.
In the agricultural sector, field gene banks have been
mainly established to provide germplasm for tropical crops,
often trees, such as cocoa, rubber, coconut, mango, cassava
and yam. The IBPGR has designated 23 field gene banks
for 9 crops, at either a global or regional level. Field gene
banks also contain wild relatives of economically important
species as well as semi-domesticated minor crops and a
number of unimproved wild plants of economic importance.
For example, the National Genetic Resources Center
(CENARGEN), Brazil, is not only the designated field gene
bank for Citrus and Arachis in Latin America, but also has
tree crops, forest trees, some vegetables and forage plants
which have recalcitrant seeds. At CENARGEN, five plants
per accession are maintained of clonal material and 50-100
seedlings of wild species are planted.
Temperate trees important for commercial forestry are
maintained in field gene banks by many national forestry
institutes and departments. These generally act as seed
orchards and for the assessment of the most suitable
genotypes for large-scale production and planting.
Many important wild tropical timber species are maintained
by tropical forestry research institutes. For example the
Arboretum de Sibang, Libreville, Gabon maintains a
collection of 40 tree taxa as a mature collection laid out in
blocks. The Forestry Research Institute, Kepong, Malaysia
maintains 722 taxa of woody species, mainly from
Southeast Asia and especially of the commercially important
dipterocarps.
Botanical gardens often have collections which are
effectively field gene banks, in that they contain significant
numbers of individuals of the same species, representing a
considerable proportion of the known wild diversity,
maintained for conservation purposes. Examples are the
native palm collection of the Jardin Botanico Nacional de
Cuba, the Universiti Kebangsaan Fernarium, Malaysia,
which has a collection of 150 out of 650 native fern and the
Lancetilla Botanic Garden and Experimental Station,
3. Conservation and Management of Biodiversity
Honduras which maintains an extensive fruit tree collection,
especially of Asiatic species (mangosteen, Citrus, mango
and rambutan), as well as 100ha of Swietenia macrophylla
(mahogany) and probably the best collection of coffee
(Coffea) cultivars in Central America.
The national collections of US endangered native plants
maintained by more than 20 botanic garden affiliates of the
Center for Plant Conservation (CPC), St Louis, have
minimum requirements for the number of individuals and
populations of a species to include and guidelines for
maximising their genetic diversity in cultivation. Over 372
species are maintained as part of the CPC national
collection.
A number of well documented natural areas of varying sizes
managed by many institutions, especially botanic gardens
and forest research institutes, function as de facto field gene
banks, combining ex situ and in situ approaches, often
through enrichmentplanting, reintroductionsand the genetic
management of indigenous plant stocks in the reserves.
There are some deficiencies with field gene banks: they
often take up a great deal of space; the collections are
generally difficult to protect from natural disasters such as
bushfires; they are susceptible to the spread of disease and
may suffer from neglect during periods of institutional
weakness. Nevertheless, for many species and in many
situations they are the only available option for the
conservation of important germplasm.
Seed banks
Seed banks are the most efficient and effective method of ex
situ conservation for sexually reproducing plants whose
seeds are suitable for long-term storage (termed orthodox
seeds). Seeds are small and therefore take up little space,
and with a few exceptions, every seed has a different
genetic constitution, so samples include a wide range of
genetic variability. At a practical level, a seed bank is
dependent on secure power supplies, the need for careful
monitoring, and testing of seed viability and the
time-consuming regeneration if the viability falls below a
certain pre-determined level and a new seed collection
cannot be made.
However it is estimated that 50,000 plant species (20% of
the world’s total) produce seeds that do not survive low
temperatures and/or dehydration. For example, many
tropical species have seeds which possess no natural
dormancy and die quickly if not allowed to germinate
immediately. These are termed recalcitrant seeds. Species
with recalcitrant seeds and those which do not readily
produce seeds need to be maintained ex situ as growing
plants in field gene banks or as living collections.
Seeds of orthodox types can be conserved for very long
periods at sub-zero temperatures if previously dried to about
5-8% moisture content. Although longevity varies from
taxon to taxon, seed viability in medium-term storage
(0-5°C) can be 5-25 years, whereas long-term storage
558
(-10°C to -20°C) gives viability of the seeds of perhaps a
hundred years.
There are many seed banks for wild plants in botanic
gardens (Table 34.11) - 528 of a total of 1,545 botanic
gardens surveyed between 1985 and 1990 have developed
a facility for seed storage and handling, with at least 144 of
them known to have low-temperature seed storage facilities.
An analysis of some selected seed bank accessions shows
the extent to which seeds of wild plants are included in
some of the world’s non-crop plant seed banks.
Some seed banks specialise in a specific geographical area
or taxonomic group. These are sometimes coordinated to
make the best use of resources, as for example in Spain
where three leading botanic institutions work in close
collaboration. The Proyecto ‘Artemis’ is a seed bank of
endemic taxa from the Iberian Peninsula and Macronesia
held at the Dep. de Biologia Vegetal, Universidad
Politécnica de Madrid, Spain. It has 1,000 of the 1,300
endemic Spanish taxa in its collections, with 1,500
accessions of known wild origin. The Jardin Botdnico de
Cordoba, Spain concentrates on the Andalucian flora of
which 300 taxa are threatened as well as 439 endemic
Iberian taxa (125 threatened) with a total of 1,498 Spanish
accessions. The Jardin Botdnico ‘Viera y Clavijo’, Gran
Canaria has in its seed bank 350 of the 500 endemic
species, most of which are threatened.
A good example of the organisation and coordination of a
local seed bank is the programme of the Centre for Plant
Conservation in the USA. This is a national network of 25
botanic gardens which together possess nearly 3,000 rare,
threatened and endangered American native species (10% of
the total American flora) as a cooperative and on a centrally
managed basis. A back-up of stored seed for plants included
in their programme is housed at the western regional station
of NPGS and at NSSL, Fort Collins, Colorado.
An important example of the development of a seed bank of
wild-collected, wild species of a crop relative is the
collection of crucifers at the Instituto Nacional de
Investigaciones Agrarias, Madrid, Spain (INIA). Over 80%
of the accessions are collected directly from the wild or in
some cases with intermediate multiplication at INIA. This
seed bank was started to conserve the wild genotypes of
Brassica and its allies.
A number of seed banks that specialise in forestry tree
species, especially ones of actual or potential economic
importance, are maintained by Forest Research Institutes
and Forestry Departments in various countries but there is
no comprehensive directory available of them or their
collections. They vary in size from regional in scope to
international.
The IBPGR and the Crop Genetic Resource Centres have
developed about 60 gene banks in the last 20 years with
long- or medium-term storage facilities of crop plants. Only
recently, however, have they included wild material and
then only of crop relatives. Wild species typically account
for less than 2% of gene bank accessions. Currently only
wild relatives of wheat (60 spp. or 75-80% of the total),
Table 34.11 Botanic garden seed banks
COUNTRY 1 2 3
ASIA
Bangladesh
China
Hong Kong
India
Indonesia
lran
Israel
Japan
Korea, Rep
Malaysia
Mongolia
Myanmar
Pakistan
Philippines
Saudi Arabia
Sri Lanka
Taiwan
Thailand
Turkey
Viet Nam
USSR (former)
USSR
EUROPE
Austria
Belgium
Bulgaria
Czechoslovakia
Denmark
Finland
France
Germany
Greece
Hungary
Ireland
Italy
Monaco
Netherlands
Norway
Poland
Portugal
Romania
Spain
Sweden
Switzerland
UK
Yugoslavia
w
ot
-~ODMDANWNHNHN WN HH HNN HK HHH UH
—-N OW —
44
as
o>}
O-W-h = =p
=
4
OONWAHN
NN
NO
=
TANONDOWAA WH ANNNWU-NYH WOOD —
wo
— =
-hwWW ao-
= =
=A
—
WBOWOUWHAWWOWN— fw OD
=
NS
Current Practices in Conservation
COUNTRY 1
NORTH AND CENTRAL AMERICA
Barbados
Belize
Canada
Costa Rica
Cuba
Dominican Republic
Grenada
Honduras
Martinique
Mexico
Panama
Puerto Rico
St Vincent
USA
SOUTH AMERICA
Argentina
Bolivia
Brazil
Chile
Colombia
French Guiana
Guyana
Paraguay
Peru
Venezuela
OCEANIA
Australia
New Zealand
AFRICA
Algeria
Angola
Benin
Cote d'Ivoire
Egypt
Ghana
Kenya
Libya
Nigeria
Réunion
Senegal
South Africa 1
Sudan
Tanzania
Togo
Tunisia
Uganda
Zimbabwe
=
AAV SSAO SAD SG yaya
ol
28
WW - KH ONNNA
=
=
(es)
NO
MND | =] | bh = @ = = = 2 2 2 a a wD
2
TOTAL 528 144 220
Source: Heywood, C.A., Heywood, V.H. and Wyse Jackson, P. 1990. International Directory of Botanical Gardens V, 5th edn. Koeltz Scientific
Books on behalf of WWF, Botanic Gardens Conservation Secretariat and International Association of Botanical Gardens.
Note: 1 Botanic gardens that report having a seed bank. 2 Botanic gardens that report having a low temperature seed storage facility. 3 Botanic
gardens that report having wild origin seeds available for distribution. Total number of botanic gardens surveyed: 1,545. Survey undertaken from
1985 to 1990.
potato (40 spp. or 70% of the total), tomato (10 spp. or
90% of the total) and to a limited extent, maize (15 spp. or
50% of the total) have been extensively collected and
preserved in seed banks.
Over 200 botanic gardens distribute seed from plants of
wild origin as part of their Index Seminum. This provides
559
a mechanism for the distribution of seeds which is more
valuable than that of unknown or garden origin, as it is
most likely to have more genetic diversity and is of known
origin. In another recent analysis of Seed Lists from botanic
gardens 432 seed lists out of 600 (from 25 countries)
included seed collected from the wild.
3. Conservation and Management of Biodiversity
In vitro Storage
Another important form of preservation of wild flora which
is carried out by many different types of institution is in
vitro storage. In vitro (literally "in glass") storage of
germplasm refers to the conservation of plants in laboratory
conditions. For germplasm storage, in vitro plants are
usually initiated from meristem tips, buds or stem tips and
propagated through division in test tubes. Jn vitro methods
are particularly suited to the long-term storage of
propagules of species with recalcitrant seeds which cannot
otherwise be maintained in a seed bank.
The plantings can be stored under various conditions but in
general at low temperatures (-3°C to -12°C) to create a
slow-growth situation and thus increase the storage period.
In vitro storage is expensive and labour-intensive, as
subculturing is necessary after a certain period (six months
to two years, depending on the species). Theoretically,
cultures can be stored indefinitely using cryogenic
techniques which would reduce labour requirements.
However, in practice only a small number of species have
yet been successfully preserved in this way, such as Malus
domestica, Ribes sp., Rubus idaeus, Vaccinium
corymboaum and Pyrus communis. More research is needed
before extensive cryobanks of wild material are established
but it is a very important development for the long-term
storage of species which are vegetatively propagated and
those with recalcitrant seeds.
Table 34.12 gives the current estimates of wild material in
tissue culture storage. The units include botanic gardens (29
units) which are the most important accounting for
approximately 1,500 taxa in vitro storage, universities (12
units) and crop research centres and private laboratories (9
units). Around 500 taxa stored in vitro worldwide are
considered threatened.
In vitro methods suffer the same disadvantages as seed
banks in terms of the need for equipment and trained staff
but techniques can be developed for local use in cooperation
with units in the developed world.
Plant reintroductions
IUCN (1987) defines reintroduction as the "intentional
movement of an organism into part of its native range from
which it is has disappeared or become extirpated as a result
of human activities or natural catastrophe". The intention is
the establishment of a self-maintaining, viable population
existing under the pressures of natural selection. The
ultimate measure of success must be the reproduction and
subsequent regeneration of the population. Reintroduction
forms one strategy aimed at the conservation of a single
species within the general umbrella of restoration that
operates at the habitat or community level.
Plant reintroductions are a high risk strategy, indications of
their long-term success are still uncertain. The intermittent
nature of plant regeneration and the ability of individuals to
survive long periods through vegetative or clonal growth
means that for woody perennials it may be many years
before regeneration is recorded.
560
Table 34.12 Wild plant material in
tissue culture storage
COUNTRY NUMBER OF UNITS NUMBER OF TAXA
China iu 3 numerous
Hong Kong 1 numerous
India 2 few
Israel 1 few
Singapore 1 few
USSR 1 few
Belgium 2 160
Denmark 2 few
France 3 few
Germany 2 few
Poland 3 few
Spain 4* 50
Sweden 3 few
UK 5 1,000
Canada 1 few
Costa Rica Usk few
Cuba 1 numerous
Mexico 1 few
USA ae numerous
Brazil 3.” numerous
Colombia hig few
Peru Us few
Australia 4* 200 +
South Africa 1 few
TOTAL 50
Source: M.F. Fay. 1991, pers. comm., Royal Botanic Gardens, Kew,
UK.
Note: Includes wild species in crop germplasm collections.
* Cryopreservation capability in one unit.
Listed below are a number of plant reintroductions
illustrating the range of plants and projects undertaken:
© Stephanomeria malheurensis (Compositae) Extinction in
the wild in 1986 due to habitat changes resulting from
alien weed invasion and associated change in fire regime
at its only known site in Oregon, USA. Seed had been
held for research, allowing subsequent reintroduction
into original site (Parenti and Guerrant, 1990).
e Pediocactus knowltonii (Cactaceae) A vulnerable
endemic restricted to a single site in New Mexico. A
joint project between the US Fish and Wildlife Service
and the State of New Mexico resulted in a second
population of 150 individuals being established in 1985
using cuttings collected from the original population.
Seed introduction has not proved successful to date, nor
has regeneration been recorded.
Sophora fernandeziana (Leguminosae) A tree native to
the Chilean islands of Juan Fernandez. The Chilean
Conservation and Forestry Organisation (CONAF) has
planted this not only to bolster low numbers of this
plant, but also because of its ecological function as a
keystone resource for the endemic hummingbird
Sephanoides fernandensis.
Ruizia cordata (Sterculiaceae) A highly endangered
shrub endemic to the Indian Ocean island of Reunion, in
April 1989 several hundred young specimens propagated
at the Brest Botanic Garden, France, were planted out on
the cliffs of the Ravine de la Grande Chaloupe, Reunion.
(Lesouef, 1991).
Gentiana nivalis (Gentianaceae) A circumpolar plant
with a restricted and diminishing distribution in Scotland.
Vulnerable to grazing, trampling and possibly climate
change. In 1980 seed was introduced to a site near to a
visitor centre in the hope of establishing a new
population that would divert damaging public attention
from the original population (Whitten, 1990).
Trochetiopsis melanoxylon (Sterculiaceae). Once thought
extinct the world population of this St Helenan endemic
is derived from only two individuals. A propagation
programme on the island has resulted in several thousand
plants being planted on the island (Drucker ef al., 1991).
A provisional survey of plant reintroductions indicates over
210 projects undertaken in over 22 countries involving 29
plant families, between 1980 and 1990. This is probably an
underestimate since plant reintroductions have traditionally
been poorly recorded and documented. The short post-
reintroduction time for these projects and the poor level of
documentation prevent an assessment of the degree of
success of these projects. A review of Californian
transplantation projects indicates a general trend: of the 15
projects reviewed 10 were unsuccessful, due to various
combinations of poor horticultural practise, poor ecological
understanding, lack of post planting maintenance and
monitoring (Hall, 1987).
Projects have been undertaken in a wide variety of habitats,
mostly in developed regions, with centres of activity in the
USA, western Europe, South Africa and Australia. Most of
the experience has been gained in temperate or
Mediterranean regions where a flora, rich in endemics,
coincides with an effective conservation infrastructure.
Botanic gardens and related institutes are holding an
increasing number of species critically threatened or extinct
in the wild, but the number of potential or required
reintroductions far exceeds the ability to undertake such
logistically demanding exercises. Because reintroductions
are long-term projects requiring extensive monitoring and
close collaboration with other agencies, they are best done
by an institute with easy access to the planting site. The
genetic viability of reintroductions originating from botanic
garden collections should be questioned as the demographic
management of cultivated plant stocks and international
co-ordination of plant conservation projects are in their
earliest stages. The material used for reintroduction comes
from a variety of sources: a species may only exist in
scattered botanic garden collections (e.g. Sophora toromiro
from Easter Island); material may be salvaged from the
existing wild populations prior to destruction of the habitat
(e.g. Penstemon barretiae from the site of a hydro-electric
project in Oregon, USA); dormant propagules may be
sampled from the soil seed bank (e.g. Jliamna corei in
Virginia, USA).
It is on the oceanic islands that reintroductions can play an
important and immediate role. This has already been
demonstrated on St. Helena and the Canary Islands; on the
former island over 8000 plants of 14 species have been
replanted (Drucker ef al., 1991). The island of Mauritius
illustrates the scale of potential reintroductions; Mauritius
has c. 112 threatened taxa with either less than 20 wild
individuals or found in 1 or 2 localities only (Owadally et
al., 1991).
561
Current Practices in Conservation
Plant reintroduction should not be regarded as a substitute
for habitat protection. In contrast it offers a technique that
can be used to upgrade the value of retained and protected
habitats. Retained areas for conservation are influenced by
increasing isolation and degradation. Accordingly
reintroduction and associated restoration programmes are
becoming accepted tools in an increasingly sophisticated
conservation regime. In the tropical nations with much
larger biological diversity and relatively poorly researched
ecology, restoration and reintroduction will play an
important future role; the work at Guanacaste, Costa Rica,
and Mineracao Rio Norte bauxite mine, Brazil, are
illustrating the potential of this work.
The scarcity of reports on past reintroductions and the need
to record and co-ordinate projects has initiated the
formation of the Reintroductions Specialist Group of the
Species Survival Commission, the Plants Group will collate
data on such projects and issue guidelines on procedure.
IN SITU CONSERVATION OF ANIMALS
Protected areas
Although it is widely accepted that protected areas are the
single most important element in the preservation of animal
species, relatively little work has been carried out to
determine how effective protected area networks are in
maintaining populations of species, either in particular
taxonomic groups or in particular geographical areas. What
work has been done, however, indicates that in many cases
a surprisingly high percentage of species are represented in
at least one protected area.
A recent study in southern Africa (Siegfried, 1989) found
that 92% of amphibian, 92% of reptilian, 97% of avian and
93% of mammalian species native to the region were
represented by breeding populations in protected areas
(Table 34.13), despite the fact that such areas covered only
6% of the land area of the region. Moreover, over 50% of
animal species were represented in more than ten reserves.
Similarly Round (1985) found that in Thailand 508 out of
578 (88%) of the native bird species were recorded from
protected areas. Of those that were not represented, 27
were mainly open country species unlikely to be adversely
affected by habitat loss and a further ten were believed
likely to occur in protected areas.
A more cursory survey of 12 African countries (Sayer and
Stuart, 1988) found that in 11 of these, at least 75%, and
generally well over 80%, of native bird species were
present in protected areas; the exception was Somalia,
where only 47% were present (Table 34.14).
Extrapolation from figures such as these indicates that in
those parts of the world which have established protected
area networks, the great majority of terrestrial species are
likely to occur in at least one, even though such areas
generally account for only a small proportion of the total
land area. Sayer and Stuart (1988) noted that just under
10% of the remaining tropical moist forest in Africa was
included in national parks and equivalent reserves, and
considered it probable that up to 90% of tropical forest
vertebrates on that continent would be maintained if these
and a few additional critical sites were adequately protected.
3. Conservation and Management of Biodiversity
Table 34.13 Breeding animal species
in protected areas in
Southern Africa
BIOME PERCENT NUMBER OF SPECIES
AMPHIBIANS REPTILES BIRDS MAMMALS
Fynbos 88.0 90.6 98.8 98.6
(22) (77) (259) (73)
Forest 100.0 100.0 99.0 100.0
(13) (21) (310) (37)
Nama-karoo 91.7 96.2 99.2 95.6
(11) (75) (250) (66)
Succulent-karoo 72.7 75.8 98.6 93.8
(8) (69) (219) (61)
Grassland 100.0 96.2 99.8 100.0
(33) (100) (416) (94)
Moist savanna 100.0 94.1 98.1 96.7
(57) (159) (530) (148)
Arid savanna 100.0 96.6 98.8 98.8
(52) (171) (513) (169)
Source: Siegfried, W.R. 1989. Preservation of species in southern
African nature reserves. In: Huntley, B.J. (Ed), Biotic Diversity In
Southern Africa. Oxford University Press, Cape Town.
Note: Absolute numbers given in parentheses.
Table 34.14 Birds in protected areas in
Africa
COUNTRY NUMBER OF SPECIES PERCENT OF
IN PROTECTED AREAS BIRD FAUNA
Cameroon 649 76.5
Céte d'Ivoire ~ 568 83.2
Ghana 558 77.4
Kenya 908 85.3
Malawi 485 777
Nigeria 719 86.5
Somalia 302 47.3
Tanzania 833 82.0
Uganda 880 89.0
Zaire 967 89.0
Zambia 637 87.5
Zimbabwe 581 S15
Source: Sayer, J.A. and Stuart, S. 1988. Biological diversity and
tropical forests. Environmental Conservation 15.
However, in.some parts of the world, including some areas
of very high diversity, the protected area network is
manifestly inadequate for the protection of a significant
proportion of the biota. This applies particularly in Oceania,
where in many countries land tenure systems make it very
difficult for significant areas of land to be set aside as
protected areas. Here, more innovative approaches to land
management (such as the Wildlife Management Areas of
Papua New Guinea) are required.
Moreover, the occurrence of a species in a protected area
is no guarantee of long-term security. Many such areas are
protected in name only and subject to continuing pressures
of encroachment, habitat degradation and hunting. Even
areas which are adequately protected are often too small to
maintain viable populations of species which live at low
population density or which are nomadic or migratory.
These problems will become increasingly pressing as
habitats outside protected areas become more and more
altered and degraded, leaving protected areas as ‘islands’ of
natural or semi-natural habitat. Under these circumstances,
the areas themselves are likely to need more active
managementif they are to maintain their ecological integrity
and continue to play their role in preventing the extinction
of species.
Recovery plans
As noted above there are circumstances in which
conventional protected areas are in themselves likely to be
inadequate for the maintenance of some animal species.
This applies when it is not possible to set aside large
enough areas to maintain viable populations of given species
or when species occur outside national boundaries (chiefly
Antarctic and pelagic organisms).
Under these conditions, recovery plans for individual
species may be developed which entail a wide range of
actions designed to improve the status of the species
concerned. The primary examples are those prepared under
the US Endangered Species Act for nationally endangered
species. Central to these is the concept of maintenance or
restoration of ‘critical habitat’, deemed to be the minimum
area of habitat necessary for the species to survive at an
acceptable level (i.e. one at which it is no longer considered
threatened). Such habitat does not necessarily have to be
within a conventional protected area, and other land-uses
may be allowed as long as they do not conflict with the
requirements of the species concerned. Frequently, different
degrees of protection are imposed on different parts of a
species’ habitat. Thus in Italy, the area of occurrence of the
Brown Bear Ursus arctos in the Apennines, centred on the
Abruzzo National Park, is divided into a variety of zones:
in the core region, no human interference is allowed, the
area being effectively a strict nature reserve devoted to the
protection of the bear and other species; surrounding this is
a region, within the park, where visitors are allowed but
human activity is strictly limited. In the area immediately
outside the park, which still constitutes important habitat for
the bear, agricultural and pastoral activities are allowed but
the bears are still strictly protected. Here compensation for
damage to livestock and crops caused by the bears is paid
in order to discourage (illegal) persecution which would
otherwise be very difficult to control. In this way,
populations of species can be maintained centred on
protected areas which would otherwise be too small to
sustain them.
As populations of individual species become smaller and
more fragmented, active management is increasingly
invoked in their conservation. This may involve
translocation of individuals from remnant, isolated
populations perceived to have no chance of long-term
survival, to larger areas of suitable habitat or to supplement
existing populations. As a last resort or as a precautionary
measure, it may also entail taking animals into captivity for
the purpose of captive breeding.
EX SITU CONSERVATION OF ANIMALS
The principal institutions holding ex situ populations of
animal species are zoos and aquaria. At least 83 countries
possess one or more zoos or aquaria (Table 34.15) but the
overall geographic distribution is very uneven: 573 (or
65%) of zoos and aquaria are located in the developed
world, Europe (298), the USA (160), Canada (24),
Australia (17), New Zealand (8) and Japan (66). These are
mainly areas of low species richness. In contrast, those
tropical regions with generally high species diversity and
large numbers of threatened species have few or no Zoos or
aquaria i.e. Africa (32), South America (29), Central
America (16), and Asia excluding China and Japan (55).
Those institutions that do exist are mainly poorly developed
and under-funded.
e
This divide is further reflected in the sizes of the zoological
collections. Institutions which reported their specimen
numbers to the International Zoo Yearbook (IZY)
collectively held approximately 1,232,000 vertebrate
specimens as of 31 December 1989 (Olney and Ellis, 1991).
Nearly half (584,000) of these were fish. The numbers of
other taxonomic groupings held were mammals (202,000
specimens), birds (351,000), reptiles (74,000) and
amphibians (21,000). The developed countries together held
67% of mammal specimens, 57% of birds, 69% of reptiles,
81% of amphibians, and 76% of fish: in total 68% of all
vertebrate specimens held.
Captive breeding - successes and shortcomings
Although most zoos have their origins as menageries for the
entertainment, and to some extent education, of the public,
they are increasingly turning their attention to conservation.
It is argued that captive populations can play a significant
role as demographic and genetic reservoirs from which
infusions of ‘new blood’ may be obtained or new
populations founded, and as last redoubts for species which
have no immediate chance of survival in the wild.
Zoos undoubtedly have considerable capacity in this regard,
but to date efforts have been relatively limited and the vast
majority of captive specimens in the world’s zoos have little
importance for the conservation of species or even in
Maintaining genetic diversity amongst non-threatened
species.
For example, although 629 mammalian species are
considered to be wholly or partly threatened on a global
scale (IUCN, 1990) only 20,628 specimens from 140
threatened species (Table 34.16) are held in zoos according
to the 1991 Census of Rare and Threatened Mammals in
Captivity (Olney and Ellis, 1991). This figure is probably
an underestimate since some collections do not respond to
the IZY’s questionnaire. In other words, although some
15% of the world’s mammal species are considered wholly
or partly threatened, only some 22% of these are
represented in captivity and only 10% of the global
zoological capacity of around 200,000 mammal specimens
consists of threatened mammal taxa. Moreover, of those
563
Current Practices in Conservation
threatened taxa, zoos included in the census make a
significant contribution to the conservation of no more than
20 full species and perhaps a similar number of subspecies.
The situation is similar (or worse) for other taxonomic
groups and is even less encouraging from the genetic
perspective. Lande and Barrowclough (1987) suggest that in
order to safeguard in the long term against the negative
genetic effects of inbreeding, a minimum viable population
of 500 individuals should be maintained. Only nine
threatened mammalian taxa have captive populations
exceeding 500 specimens, and only a further 14 have
captive populations exceeding 250 (Table 34.16).
A criticism which is often made of captive breeding
programmes is that they are a misallocation of resources.
The basis for this criticism is that large amounts of money
are spent on captive breeding efforts in comparison with
that available for in situ conservation despite the fact that
captive breeding is much less cost-effective than
preservation in situ. For example, Leader-Williams (1990)
calculates that the cost of keeping African elephants and
black rhinos in zoos is 50 times that of protecting equivalent
numbers in the wild in Zambian National Parks, where
1km? of park can be adequately patrolled for the annual sum
of only US$400. In addition, the maintenance of captive
populations does not have the associated benefits of
protecting an organism’s habitat, and by logical extension
thousands of other species.
Zoos are sensitive to these criticisms, and a significant
number are attempting to improve their efforts in the
conservation of threatened species, chiefly through
improved international cooperation and clearer setting of
priorities for breeding threatened species, as well as by
devoting larger fractions of their budgets to field
conservation. Coordination in these efforts is carried out
through a variety of interconnected mechanisms, including
studbooks, the TUCN/SSC Captive Breeding Specialist
Group (CBSG), the International Species Inventory System
(SIS) and a number of regional cooperative captive
breeding programmes.
Studbooks
In order to facilitate the success of captive-breeding
programmes (e.g. to help prevent inbreeding) and to aid in
the development of successful management techniques a
series of studbooks have been developed. A studbook is an
international register which lists all captive individuals of a
taxon of conservation concern. Official studbooks are those
recognised by the Species Survival Commission (SSC) of
IUCN and the International Union of Directors of
Zoological Gardens. They are coordinated through the
International Zoo Yearbook and the CBSG. As of August
1991 there were 104 recognised International Studbooks and
five International Registers (1 amphibian, 4 reptiles, 19
birds and 85 mammals, see Table 34.17), with a further
three studbook applications awaiting endorsement (Olney
1991). In principle, studbooks are published every three
years, and regular updates are available.
3. Conservation and Management of Biodiversity
Table 34.15 Number of vertebrates held in zoos and aquaria
NUMBER OF NUMBER OF VERTEBRATES HELD
ZOOS AND Mammals Birds Reptiles Amphibians Fishes TOTAL
AQUARIA
WORLD 878 201706 351484 T4416 20788 583832 1232226
ASIA 252 46646 110121 15957 3380 281878 457982
Bahrain 1 575 600 16 18 60 1269
Brunei 1 () 0 7 ts) 492 499
China 131 12489 46175 1348 1121 70406 131539
- Hong Kong 2 236 3002 220 te) 2680 6138
India 17 5045 10605 5148 0 2607 23405
Indonesia 4 1847 3683 717 58 1808 8113
Israel 4 1721 1977 964 90 () 4752
Japan 66 12554 17653 4503 2014 195530 232254
Korea, Rep 4 1903 6070 130 (0) 261 8364
Kuwait 1 208 493 34 0 i) 735
Malaysia 2 663 944 302 ts) 2336 4245
Myanmar 1 968 599 492 0 i) 2059
Pakistan 2 = = = = = =
Philippines 1 = = = = = =
Qatar 2 900 401 71 4 (e) 1376
Saudi Arabia 1 700 18 te) te) 0 718
Singapore 3 689 4662 279 18 1600 7248
Sri Lanka 1 741 1282 510 ie) 1946 4479
Taiwan 1 1118 1221 113 to) ts) 2452
Thailand 4 2295 7692 475 te) 387 10849
Turkey 1 139 565 15 te) 649 1368
United Arab Emirates 2 1855 2479 613 57 1116 6120
USSR (former)
40 16981 22557 3544 1714 30735 75531
EUROPE: 298 72558 115933 21936 8949 129463 348839
Austria 5 1065 1158 603 204 2108 5138
Belgium 5 2409 2280 421 105 5531 10746
Bulgaria 2 274 490 67 1429 ) 2260
Czechoslovakia 15 4311 4875 701 30 4437 14354
Denmark 7 1770 1593 424 278 4037 8102
Finland 3 727 602 36 6 16 1387
France 34 6520 11226 1538 34 428 19746
Germany 56 21850 35163 6667 2482 31112 97274
Hun: 6 1409 1645 214 9 2084 5361
Ireland 2 747 770 30 1 te) 1548
Italy 21 2058 3545 578 382 2525 9088
Monaco 1 = = = = = =
Netherlands 11 4365 7717 2287 1389 8578 24336
Norway 1 10 22 (0) 9 1500 1541
Poland 9 3290 3547 1659 325 8891 17712
Portugal 3 593 1400 258 (0) 3727 5978
Romania 1 297 572 28 () () 897
Spain 11 2106 7185 888 42 6073 16294
Sweden 12 1775 1573 592 403 6222 10565
Switzerland 15 2625 4520 849 314 5158 13466
United Kingdom 76 14200 25779 4014 1507 19036 64536
Yugoslavia 2 157 271 82 0 18000 18510
NORTH AND CENTRAL AMERICA 20t 47713 65299 23700 6078 115706 258586.
Barbados 1 236 125 111 20 26 518
Belize 1 55 45 72 () ty) 172
Bermuda 1 16 198 55 t) 951 1220
Canada 24 4546 4511 1632 370 9928 20987
Cuba 4 430 1178 195 () 90 1893
Dominican Republic 1 - - - - - -
Jamaica 1 = = = = = >
Mexico 5 2637 7605 587 (e) 167 10996
Netherlands Antilles 1 - = - = - =
Puerto Rico 1 135 272 88 () 20 515
Trinidad and Tobago 1 165 403 99 310 851 1828
United States 160 39493 50962 20861 5378 103763 220457
SOUTH AMERICA 2 $403 13060 4064 67 9240 31834
Argentina 3 595 1500 329 24 150 2598
Bolivia 1 358 1409 118 24 () 1909
Brazil 10 2368 6473 2054 te) 550 11445
Chile 1
Colombia 2 407 787 324 i*} i) 1518
Guyana 1 70 93 43 te) te) 206
Peru 1 385 346 118 2 te) 851
Uruguay 3 596 762 185 9 0 1552
Venezuela 7 624 1690 893 8 8540 11755
OCEANIA 26 6035 9883 2244 210 MOEA 22456
Australia 17 5181 7944 1815 163 3766 18869
New Zealand 8 854 1939 429 47 318 3587
Papua New Guinea 1 = - - - - =
AFRICA 32 6370 14631 2971 390 12636 36008
Egypt 2 256 2317 36 11 0 2620
Ghana 1 52 76 55 te) 0 183
Kenya 4 1562 187 14 0 0 1763
Libya 1 555 613 172 te) () 1340
Madagascar 2 t') () 43 ) () 43
564
Table 34.15
Current Practices in Conservation
Number of vertebrates held in zoos and aquaria (contined)
NUMBER OF NUMBER OF VERTEBRATES HELD
ZOOS AND Mammals Birds Reptiles Amphibians Fishes TOTAL
AQUARIA
AFRICA (continued)
Mauritius 1 = = = = = -
Morocco 1 672 1213 28 ie) 0 1913
Nigeria 3 130 130 118 0 ie) 378
Senegal 1 = = ~ = = =
South Africa 12 2611 9237 1736 379 12636 26599
Sudan 1 = = = = = =
Tunisia 1 332 658 740 is} 0 1730
Zaire 1 = ~ = = = =
Zimbabwe 1 200 200 29 ts) ts) 429
Source: Olney, P-J.S. and Ellis, P. (Eds) 1991. 1990 International Zoo Yearbook, Vol.30. Zoological Society of London, London.
Note: Some institutions did not report specimen numbers.
International Species Inventory System
Since 1974 these studbooks have been supplemented by the
International Species Inventory System (ISIS), a global
information network designed to support sound genetic and
demographic management of zoological collections and
enable zoos to meet their increasing conservation
responsibilities. ISIS maintains a centralised computer
database of census, demographic, genealogical, and
laboratory data on wild animals held in captivity. In
September 1991 the database contained information on over
141,480 living vertebrate specimens from 4,200 taxa, held
in more than 395 zoological institutions in 39 countries,
plus an even greater number of their ancestors. ISIS has
very good coverage of North American zoos, coverage of
Europe and Australasia is rapidly expanding, while
participation by institutions in Latin America, Asia and
Africa should be increased.
All participating institutions receive a 15,000+ page ISIS
Species Distribution Report on microfiche every six
months, and copies of annual bound ‘Abstracts’, one each
for mammals, birds, reptiles and amphibians. These include
information on sex and age distribution, births, deaths, and
important trends for the species as a whole. Although
historically zoos were able to draw on populations of wild
animals to supply their specimens, this is often no longer
feasible because of ethical and ecological considerations.
Instead, greater reliance is placed on captive breeding. ISIS
data indicate that 92% of new zoo mammals are now
captive-bred, along with 71% of birds and a majority of
reptiles and amphibians (Anon., 1991a).
By 1990, these various programmes had resulted in over
150 taxa of all classes of vertebrates as well as the
invertebrate genus Partula (endemic land snails of Moorea,
now extinct in the wild) being managed cooperatively by
groups which totalled about 400 institutions - approximately
half the world’s zoos (Flesness and Foose, 1990). Zoos
plan to expand the number of species now involved in
multi-institution breeding programmes from 150 to 1,000
(Flesness and Foose, 1990).
Animal reintroduction programmes
One area where international networks and coordination of
the world’s zoos can play a role in in situ conservation is
through reintroduction programmes. The artificial
movement of individual animals between populations is
becoming increasingly used as a conservation tool. Griffith
565
et al. (1989) reported that over 700 translocations or
repatriations occurred each year, mainly in the USA and
Canada. These projects are frequently conducted with the
support of international captive-breeding programmes at
zoological gardens and aquaria, and may generate much
public enthusiasm.
Reintroduction projects are not always successful. Griffith
et al. (1989) examined the outcome of projects involving
birds and mammals. Native game species constituted 90%
of translocations and had a higher success rate (86% of 118
projects considered) than translocations of threatened,
endangered or sensitive species (44% of 80 projects). Dodd
and Siegel (1991) found an even lower overall success rate
of only 19% for 25 projects involving reptiles and
amphibians.
The reasons behind the high failure rates of reintroduction
attempts are diverse. Most importantly, not all species lend
themselves to reintroduction. In addition a variety of
ecological factors can affect the success of reintroduction
programmes, including the quality of the habitat in which
the release occurs, whether the individuals released are wild
or captive bred, and the feeding habits of adults. The design
of the reintroduction is also crucial: such factors as the
number, sex and social composition of individuals released,
whether the release is ‘hard’ (no food and shelter provided
on site) or ‘soft’, and planning for further releases after
populations become established in order to provide
injections of new blood, may all be important. In order to
succeed reintroduction must therefore be carefully planned
and executed, and monitoring should ideally continue for
several generations after release.
Case study: reintroduction of Arabian Oryx to Oman
The Arabian (or White) Oryx, Oryx leucoryx formerly
inhabited arid gravel plains and sandy deserts throughout
the Arabian Peninsula and adjacent regions. By the early
1960s the species was confined to two small areas: where
the borders of Saudi Arabia, Yemen and Oman meet; and
in north-eastern Oman. The last wild oryx were probably
killed in 1972 in the Jiddat al-Harasis of Oman (Henderson,
1974) although rumours of sightings persist.
Fortunately, significant numbers of oryx remained in
captivity in the Middle East and elsewhere, notably at
Phoenix in Arizona, USA, where a herd had been
established in the 1960s in response to the continued
depletion of the species in the wild. In 1974 the “White
3. Conservation and Management of Biodiversity
Table 34.16 Census of IUCN threatened animals held in captivity
IUCN NUMBERS HELD IN CAPTIVITY NUMBERS OF CAPTIVE ANIMALS
THREAT BRED IN CAPTIVITY
CATEGORY MALE FEMALE UNKNOWN TOTAL MALE FEMALE UNKNOWN TOTAL
MAMMALS 20770 5500
Zaglossus bruijni Vv 2 5 0 if 0 1 ts) 1
Myrmecobius fasciatus E 5 6 (0) 11 2 3 ts) 5
crotis lagotis E 16 11 0 27 16 11 it) 27
Gymnobelideus leadbeateri Vv 23 15 3 41 20 13 6 39
Potorous longipes | 7 4 i) 11 6 3 () 9
Bettongia penicillata E 90 97 6 193 t) ts) tt) Most
Solenodon paradoxus |= 1 ty) 0 1 te) 0 0 0
Pteropus rodricenss E 64 70 18 152 0 i) 0 Most
Macroderma gigas Vv 4 9 0 13 3 6 0 9
Microcebus coquereli Vv 31 29 4 64 () () ) Most
Lemur coronatus E 20 17 0) 37 16 13 (0) 29
Lemur m. macaco Vv 109 97 4 210 t) tt) te) Most
L. m. albifrons R 96 79 8 183 ts) te) ts) Most
L. m. collaris Vv 19 18 0 37 15 16 Oo 31
L. m. flavirons E 9 4 te) 13 6 1 () 7
L. m. fulvus R 60 65 25 150 (0) ts) () Most
L. m. mayottensis Vv 49 54 18 121 ts) t') (1) Most
L. m. rufus R 33 42 3 78 ts) ts) () Most
L. m. santordi Vv 9 10 () 19 7 8 tt) 15
Lemur mongoz E 38 34 () 72 () () 0 Most
Lemur rubriventer Vv 5 5 te) 10 2 2 () 4
Hapalemur griseus K 9 10 te) 19 6 5 te) 11
Varecia variegata E 226 250 8 484 (°) te) (0) Most
Propithecus fattersalli E 1 2 te) 3 t) 1 ts) 1
Propithecus verreauxi Vv 6 8 (*) 14 2 3 0 5
Daubentonia madagascariensis E 2 1 tt) 3 te) ts) t) t)
Tarsius syrichta E 10 13 2 25 2 5 5 12
Callithrix aurita E 5 3 1 9 4 3 1 8
Callithrix humeralfer K 5 5 2 12 3 4 2 9
Callithrix jacchus flaviceps IS 1 0 (e) 1 0 () (e) ()
Saguinus bicolor E 16 18 9 43 13 8 9 30
Saguinus imperator u 96 82 8 186 0 0 te) Most
Saguinus o. oedipus E 561 509 95 1165 0 tt) tt) Most
Leontopithecus r.rosala E 274 285 33 592 0 te) tt) Most
L. chrysomeks E 109 86 29 224 83 64 28 175
L. chrysopygus E 33 28 8 69 23 20 14 57
Callicebus personatus = 2 0 {0} 2 (0) 0 () 0
Callimico goeldii R 156 140 21 317 te) 0 t) Most
Saimiri oerstedi E 2 2 () 4 te) 1 tt) 1
Chiropotes albinasus Vv i) 2 i} 2 0 2 0 2
Chiropotes s. satanas = 0 2 0 2 (0) [) 0 )
Cacajao c. calvus Vv 1 1 (+) 2 0 ) o 0
C. c. rubicundus Vv 3 5 0 8 2 2 t) 4
Cacajao melanocephalus Vv 2 1 (0) 3 0 0 0 0
Alouatta fusca (=guariba) Vv 5 0 (0) 5 tt) 0 it} te)
Ateles belzebuth Vv 31 56 5 92 14 20 4 38
Ateles fusciceps Vv 29 46 (0) 75 20 21 it) 41
Ateles paniscus Vv 81 137 2 220 46 59 1 106
Brachyteles arachnoides E 1 3 0 4 0 0 0 0
Lagothrix lagothricha Vv 49 62 () 141 22 42 te) 64
Macaca silenus E 188 207 7 402 0 0 ts) Most
Macaca sylvanus Vv 368 483 23 874 i) 0 0 )
Cercocebus t. torquatus Vv 60 46 (0) 106 48 30 ° 78
C. t. atys Vv 73 140 te) 213 65 121 te) 186
C. t. lunulatus Vv 11 17 () 28 7 10 () 17
Mancrillus leucophaeus E 22 30 0 52 0 (0) 0 0
Theropithecus gelada R 40 70 0 110 0 ty) 0 0
Cercopithecus diana Vv 86 106 0 192 63 63 5 131
Cercopithecus hamlyni Vv 20 30 2 52 13 20 2 35
Cercopithecus lhoesti Vv 6 5 1 12 5 2 1 8
Cercopithecus preussi E () 1 (0) 1 te) t) tt) tt)
Allenopithecus nigroviridis K 28 27 4 59 8 10 4 22
Pygathrix nemaeus E 25 29 tt) 54 14 16 tt) 30
Nasalis larvatus Vv 8 10 (¢) 18 6 7 te) 13
Presbytis francoisi E 19 33 (°) 52 10 12 te) 22
Presbytis geei R 1 9 17 37 (0) 2 () 2
Presbytis johni E 10 10 6 26 6 1 0 7
cma concolor Vv 67 66 6 139 30 18 5 53
lylobates klossi E 2 2 0 4 (0) te) te) ts)
Hylobates moloch E 15 12 1 28 6 4 1 1
Hylobates pileatus E 33 32 0 65 5 2 0 7
Pongo pygmaeus E 299 379 2 680 205 255 2 462
Pan paniscus Vv 30 36 ) 66 20 19 0 39
Gorilla g. gorilla Vv 278 339 0 617 146 154 () 300
G. g. graueri E 3 3 () 6 () 2 (*) 2
Myrmecophaga tridactyla Vv 47 51 2 100 18 17 1 36
Romerolagus diazi |= 3 1 ts) 4 3 1 0 4
Geocapromys browni R 4 3 ts) 7 4 3 0 a
Plagiodontia aedium R 1 2 0 3 0 1 0 1
Chrysocyon brachyurus Vv 132 127 2 261 (*) 0 0 Most
Speothos venaticus Vv 42 44 2 88 0 i) 0 Most
Cuon alpinus Vv 29 28 0 57 18 20 tt) 38
Lycaon pictus E 146 123 0 269 0 0 i) t)
Tremarctos ornatus Vv 58 64 4 126 43 45 4 92
Melursus ursinus Vv 50 52 4 106 26 23 1 50
Ailuropoda melanoleuca E 6 2 1 9 2 0 1 3
Lutra |. longicaudis (+ platensis) Vv 7 5 0 12 0 ts) 0 i)
Lutra |. luta Vv 56 99 5 160 33 70 5 108
Pteronura brasiliensis Vv 8 6 0 14 0 (0) ts) tt)
Cryptoprocta ferox K 13 8 t) 21 1 4 (e) 15
566
Current Practices in Conservation
Table 34.16 Census of IUCN threatened animals held in captivity (continued)
IUCN NUMBERS HELD IN CAPTIVITY NUMBERS OF CAPTIVE ANIMALS
THREAT BRED IN CAPTIVITY
CATEGORY MALE FEMALE UNKNOWN TOTAL MALE FEMALE UNKNOWN TOTAL
MAMMALS (continued)
Hyaena brunnea Vv 16 12 0 28 9 6 0 15
Felis margarita scheffeli E 2 1 t) 3 2 1 t) 3
Felis marmorata | 2 4 0 6 1 3 0 4
Felis planiceps I 1 ts) 0 1 0 0 0 0
Felis temmincki | 28 20 2 50 () 0 ) Most
Felis tigrina Vv 10 7 0 17 1 1 (0) 2
Felis wiedi Vv 39 41 1 81 12 20 1 33
Panthera leo persica E 68 80 0 148 is) is) 0 Most
Panthera pardus delacouri Lf 4 4 0 8 3 4 0 if
P. p. fusca if 10 14 0 24 9 14 0 23
P. p. japonensis T 31 29 0 60 0 0 0 Most
P. p. kotiya T 28 21 0 49 8 8 0 16
P. p. orientalis il 39 28 1 68 (0) 0 0 Most
P. p. saxicolor 1 61 68 0 129 is) (9) 0 Most
Panthera t. tigris E 72 75 0 147 () () () Most
P. t. corbetti E 5 6 te) 11 1 5 ts) 6
P. t. altaica [2 325 387 te) 712 te) 0 ts) Most
P. t. amoyensis E 33 19 0) 52 1) (0) 0 Most
P.t. sumatrae E 7 88 te) 165 ts) t) t) Most
Panthera uncia E 185 182 ft) 367 Qo () L*) Most
Neofelis nebulosa Vv 91 75 6 172 68 59 2 129
Acinonyx jubatus Vv 250 275 0 525 175 170 ts) 345
Dugong dugon Vv 1 1 0 2 i) 0 0 0
Trichechus inunguis Vv 1 0 0 1 0 (0) 0 (0)
Trichechus manatus Vv 12 12 () 24 6 1 0 7
Equus przewalskii Ex? 397 564 0 961 397 564 te) 961
Equus hemionus onager Vv 42 70 0 112 ts) te) te) Most
E. h. kulan Vv 93 191 0 284 0 0 0 Most
E. h. khur E 4 7 0 ant 2 2 0 4
E. h. kiang Vv 30 25 te) 55 30 25 () 55
Equus africanus E 9 6 0 15 7 3 (0) 10
Equus grevyi E 135 250 0 385 119 188 0 307
Equus zebra hartmannae Vv 40 79 0 119 40 79 0 119
Tapirus bairdi Vv 15 14 0 29 11 8 0 19
Tapirus indicus E 58 72 1 131 41 54 1 96
Tapirus pinchaque Vv 5 4 0 9 1 2 0 3
Rhinoceros unicornis E 55 40 14 109 32 21 1 54
Dicerorhinus sumatrensis E 4 9 0 13 1 ts) 0 1
Ceratotherium s. simum E 4 5 (0) 9 2 3 0 5
Diceros bicornis E 61 83 (¢) 144 35 45 () 80
Babyrousa babyrussa Vv 34 21 3 58 (0) te) 0 Most
Choeropsis liberiensis Vv 81 115 te) 196 63 87 0) 150
Vicugna vicugna Vv 61 62 ts) 123 ts) 0 0 Most
Muntiacus feaei E 10 9 (0) 19 5 2 0 7
Dama dama mesopotamica E 11 21 (0) 32 () tt) te) Most
Axis calamianensis Vv 3 3 0 6 3 1 0 4
Axis kuhli R 14 21 2 37 Mi 8 2 il?
Cervus duvauceli E 101 194 10 305 te) 0 0 Most
Cervus elaphus bactrianus E 16 20 (0) 36 0 0) (0) Most
Cervus e. eldi Vv 22 22 6 50 (0) (e) (e) Most
C. e. siamensis EB 1 2 (0) 3 (0) (e) 0 0
Cervus nippon taiouanus E 104 182 32 318 104 182 32 318
Blastocerus dichotomus Vv 3 10 (0) 13 3 5 0 8
Taurotragus derbianus E 13 16 ie) 29 10 11 ts) 2i
Bubalus (Anoa) depressicornis E 23 20 0 43 ie) () te) Most
Bubalus (Anoa) quarlesi = 5 3 0 8 3 2 0 5
Bos gaurus Vv 80 125 te) 205 te) 0 0 Most
Bos javanicus Vv 79 133 1 213 i) 0 0 Most
Cephalophus jentinki E 5 4 0 9 4 3 0 7
Cephalophus zebra Vv 3 2 () 5 1 1 i) 2
Kobus leche Vv 105 236 2 343 0 () (0) Most
Oryx dammah E 225 424 5 654 0 () ts) Most
Oryx leucoryx E 209 269 13 491 (0) te) 0 Most
Addax nasomaculatus E 179 310 7 496 ts) te) ts) Most
Damaliscus d. dorcas Vv 25 54 () 79 (e) 0 0 Most
Damaliscus hunteri Vv 2 1 0 3 1 1 0 2
Gazella cuvieri E 33 72 ie) 105 0 0 tt) Most
Gazella dama E 40 82 0 122 (0) 0 0 Most
Gazella d. mhorr E 56 86 (0) 142 te) 0 0 All
Gazella dorcas isabella Vv 3 17 1 21 3 16 1 20
G. d. massaesyla Vv 2 2 0 4 2 2 0 4
G. d. osiris (incl. neglecta) Vv 42 88 0 130 (0) (0) 0 Most
G. d. saudiya Vv 18 40 2 60 17 37 2 56
Gazella lla arabica Vv 74 105 18 197 0 (0) 0 Most
Gazella leptoceros E 48 71 (0) 119 (0) ie) 0 Most
Gazella rufifrons Vv 0 2 is) 2 (0) (e) 0 ts)
Gazella spekei Vv 15 21 to) 36 (0) (0) 0 Most
Gazella subgutturosa marica E 165 185 7 357 (0) 0 ts) Most
Rupicapra rupicapra ornata Vv 3 13 5 21 3 11 5 19
Hemitragus hylocrius Vv 12 25 0 37 12 25 () 37
Capra falconeri Vv 45 60 4 109 0 0 0 Most
C. f. megaceros (incl. jerdoni) E 7 15 0 22 0 0 0 Most
BIRDS 16736 9487
Apteryx oweni Vv 2 2 1 5 te) 1 1 2
Ciconia boyciana R 10 8 te) 18 3 1 0 4
Geronticus eremita E 145 157 334 636 0 0 0 Most
Cairina scutulata Vv 79 64 4 147 ie) 0 0 Most
Anas aucklandica chorotis R 18 21 32 71 te) () () Most
567
3. Conservation and Management of Biodiversity
Table 34.16 Census of IUCN threatened animals held in captivity (continued)
IUCN NUMBERS HELD IN CAPTIVITY NUMBERS OF CAPTIVE ANIMALS
THREAT BRED IN CAPTIVITY
CATEGORY MALE FEMALE UNKNOWN TOTAL MALE FEMALE UNKNOWN TOTAL
BIRDS (continued) 13775 8824
Gymnogyps californianus E 7 9 te) 16 1 2 Uy) 3
Haliaeetus albicilla R 63 77 45 185 13 20 11 a
Haliaeetus pelagicus R 12 12 7 31 2 1 0 3
Harpia harpyja R 21 14 3 38 5 ts) ts) 5
Macrocephalon maleo Vv 1 3 2 6 2 () () 2
Crax mitu E 30 27 8 65 te) ts) 0 Most
Crax blumenbachi E 4 4 tt) 8 3 3 (0) 6
Lophura bulweri R 15 15 0 30 tt) 0 0 Most
Grus nigricollis R 2 2 tt) 4 0 0 0 ts)
Gus monacha R 30 47 10 87 16 31 5 52
Gus japonenss Vv 113 126 28 267 76 90 14 180
Gus americana E 23 24 8 55 16 22 8 46
Gus vipio R 102 107 49 258 67 74 30 171
Grus leucogeranus R 25 20 0 45 8 7 te) 15
Rhynochetos jubatus E 1 1 0 2 0 () 0 0
Nesoenas mayeri E 37 32 26 95 0 te) ts) Most
Psephotus c. chrysopterygius R 29 25 is) 54 te) te) 0 Most
Aratinga guarouba Vv 76 66 24 166 32 25 3 60
Rhynchopsitta pachyrhyncha Vv 50 50 18 118 17 14 11 42
Amazona pretrei Vv 15 9 (0) 24 (°) 1 0 1
Amazona versicolor R 5 6 11 22 1 2 8 11
Amazona arausiaca E 1 1 te) 2 (0) te) i) (0)
Amazona guildingi R 15 16 1 32 4 4 i) 8
Pharomachrus mocinno Vv 1 1 0 2 te) 0 0 te)
Picathartes gymnocephalus Vv 1 2 0 3 (°) 1 i) 1
Picathartes oreas Vv 0 1 0 1 0 1 0 1
Foudia flavicans E 2 4 0 6 2 4 (0) 6
Leucopsar rothschildi e 174 177 111 462 0 0 ts) Most
REPTILES 6187 4279
Batagur baska E 8 3 1 12 0 0 0 0
Clemmys muhlenbergi R 26 26 11 63 12 9 11 32
Terrapene coahuilla Vv 10 12 95 117 7 9 95 111
Geochone elephantopus Vv 77 51 108 236 13 15 107 135
Geochelone radiata Vv 86 60 321 467 7 10 251 268
Geochelone yniphora E 6 8 21 35 0 1 21 22
Pyxis arachnoides | 4 2 te) 6 t) 0 te) t)
Pseudoemydura umbrina E 6 3 6 15 3 1 6 10
Alligator sinensis E 20 29 31 80 8 22 27 57
Caiman latirostris [= 70 30 166 266 11 3 48 62
Melanosuchus niger E 4 4 8 16 0 0 0 0
Crocodylus intermedium E 2 3 6 11 0 1 () 1
Crocodylus moreleti E 15 17 27 59 11 15 17 43
Crocodylus palustris Vv 455 1881 728 3064 414 1852 647 2913
Crocodylus rhombifer E 22 34 25 81 12 8 20 40
Gavialis gangeticus E 21 43 192 256 3 13 157 173
Sphenodon punctatus R 15 14 34 63 1 i) 24 25
Phelsuma guentheri E 10 24 3 37 0 0 i) Most
Brachylophus fasciatus Vv 20 19 7 46 to) i) 0 Most
Leiolopisma tefairi R 7 9 2 18 (°) () i} Most
Heldoderma horridum | 25 14 35 74 3 te) 16 19
Heloderma suspectum Vv 62 55 73 190 13 9 19 41
Varanus griseus caspius Vv 18 11 13 42 ie} 0 ie) 0
Varanus komodoensis R 2 6 4 12 2 0 0 2
Epicrates angulifer | 63 64 49 176 34 34 47 115
Epicrates inornatus E 27 24 34 85 17 12 31 60
Epicrates striatus fosteri R 2 te) fe) 2 te) 0 ) te)
Epicrates subflavus Vv 57 38 12 107 39 32 7 78
Acrantophis dumerik K 77 95 3 175 it) 0 0 Most
Acrantophis madagascariensis K 19 27 4 50 8 17 3 28
Sanzinia madagascariensis K 51 44 13 108 to) t°) 0 Most
Naja oxiana [= 19 15 10 44 2 5 3 10
Vipera lebetina schweizeri Vv 3 3 (0) 6 3 2 i) 5
Vipera raddei i 14 20 9 43 5 6 4 15
Crotalus unicolor R 37 39 1 77 () 0 (0) Most
Crotalus willardi iT 28 14 6 48 10 2 2 14
AMPHIBIANS 206 68
Andrias davidianus | 2 2 49 53 (0) te) (0) 0
Andtias japonicus R 12 8 119 139 68 te) ie) 68
Typhlomolge rathbuni E 6 0 (0) 6 (0) 0 0 0
Bufo houstonensis E 2 2 0 4 ts) 0 te) ts)
Conraua goliath Vv 4 0 0 4 0 ce) ie) i)
Sources: data from Olney, PJ.S. and Ellis, P. (Eds) 1991. 1990 International Zoo Yearbook, Vol.30. Zoological Society of London, London. [UCN
Threat Categories obtained from IUCN 1990. 1990 IUCN Red List of Threatened Animals. 1UCN, Gland.
Notes: It should be noted that in some cases the taxonomy of the International Zoo Yearbook differs from that of the IUCN Red List. The numbers given
represent minimum estimates of species numbers held in captivity since private collections and other non—zoo organisations fall outside the scope of the
International Zoo Yearbook. A number of species considered threatened by IUCN in their natural habitats are relatively common in captivity and breed
regularly. For reasons of space, these taxa (which include chimpanzee, wolf, polar bear, jaguar, and various species of ducks, pheasants and reptiles) were
See omitted aboeether rom the International Zoo Yearbook census, or only their total captive populations have been registered. No data were available
or threatened fish.
568
Current Practices in Conservation
Table 34.17 Current Studbooks and International Registers
Amphibians
Reptiles
Birds
Mammals
SPECIES
Bufo /emur
Alligator sinensis
Heloderma suspectum
Heloderma horridum
Crotalus unicolor
Apteryx spp.
Geronticus eremita
Tragopan blythi
Tragopan caboti
Lophura edwardsi
Crossoptilon crossoptilon
Polyplectron inopinatum
Polyplectron malacense
Afropavo congensis
Grus monacha
Grus japonensis
Grus vipio
Grus leucogeranus
Bugeranus carunculatus
Colomba (Nesoenas) mayeri
Cyanopsitta spixii
Aratinga gauroba
Amazona guildingii
Buceros bicornis
Bettongia penicillata
Dendrolagus matschiei
Lemur m. macao
Lemur mongoz
Varecia variegata
Nycticebus pygmaeus
Cebuella pygmaea
Saguinus imperator
Saguinus o. oedipus
Leontopithecus r. rosalia
Leontopithecus chrysomelas
Leontopithecus chrysopygus
Callimico goeldii
Alouatta caraya
Macaca silenus
Mandrillus leucophaeus
Theropithecus gelada
Cercopithecus d. diana
Pygathrix nemaeus
Hylobates concolor
Hylobates moloch
Hylobates pileatus
Pongo pygmaeus
Pan paniscus
Gorilla gorilla
Myrmecophaga tridactyla
Dinomys branicki
Canis lupus baileyi
Canis rufus
Chrysocyon brachyurus
Speothos venaticus
Tremarctos ornatus
Ursus maritimus
Ailurus fulgens
Ailuropoda melanoleuca
COMMON NAME
Puerto Rican Crested Toad
Chinese Alligator
Gila Monster
Beaded Lizard
Aruba Island Rattlesnake
Kiwis
Red-cheeked Ibis (Waldrapp)
Blyth’s Tragopan
Cabot’s Tragopan
Edward’s Pheasant
White Eared Pheasant
Rothschild’s Peacock
Malayan Peacock Pheasant
Congo Peafowl
Hooded Crane
Red-crowned Crane
White-naped Crane
Siberian White Crane
Wattled Crane
Mauritius Pink Pigeon
Spix’s Macaw
Golden (Queen of Bavaria)
St Vincent Parrot
Great Indian Hornbill
Brush-tailed Bettong
Matschie’s Tree Kangaroo
Black Lemur
Mongoose Lemur
Ruffed Lemur
Pygmy Loris
Pygmy Marmoset
Emperor Tamarin
Cottontop Tamarin
Golden Lion Tamarin
Golden-headed Lion Tamarin
Black Lion Tamarin
Goeldi’s Monkey
Black Howler Monkey
Lion-tailed Macaque
Drill
Gelada Baboon
Diana Monkey
Douc Langur
Black Gibbon
Moloch Gibbon
Pileated Gibbon
Orang-Utan
Bonobo
Gorilla
Giant Anteater
Pacarana
Mexican Wolf
Red Wolf
Maned Wolf
Bush Dog
Spectacled Bear
Polar Bear
Lesser or Red Panda
Giant Panda
569
IUCN STATUS
CATEGORY
—-DV—-m<DvD<—m
DommMmaAaADW<K DAD:
m
mammmm—-: cmmee
maAc<c<ccmemeccmmmemeanmm
*
*
STUDBOOK (S)
OR REGISTER (R)
®OAPYD®BDADDODADDADDDADDDHDUHDHHDHDHAVDHHHHHAHHHHDDHHHHAHHHHAHHHHHHHHHRDHNHDHHHHVHYDY
3. Conservation and Management of Biodiversity
Table 34.17 Current Studbooks and International Registers (continued)
SPECIES COMMON NAME IUCN STATUS STUDBOOK (S)
CATEGORY OR REGISTER (R)
Mammals Lutra /. lutra European Otter Vv Ss
(continued) Aonyx cinerea Oriental Small-clawed Otter K Ss
Hyaena brunnea Brown Hyena v Ss
Felis margarita Sand Cat - Ss
Felis nigripes Black-footed Cat - Ss
Panthera leo persica Asiatic Lion E Ss
Panthere pardus sspp? Leopard i Ss
Panthera tigris sspp* Tiger E Ss
Panthera uncia Snow Leopard E Ss
Neofelis nebulosa Clouded Leopard Vv Ss
Acinonyx jubatus Cheetah Vv Ss
Equus przewalskii Przewalski’s Horse Ex? Ss
Equus hemionus Asiatic Wild Ass Vv Ss
Equus africanus African Wild Ass E Ss
Equus grevyi Grevy’s Zebra E Ss
Equus zebra hartmannae Hartmann’s Zebra Vv Ss
Tapirus bairdi Baird’s Tapir Vv Ss
Tapirus indicus Malayan Tapir E =)
Rhinoceros unicornis Indian Rhinoceros E Ss
Dicerorhinus sumatrensis Sumatran Rhinoceros E Ss
Ceratotherium simum White Rhinoceros - Ss
Diceros bicornis Black Rhinoceros E Ss
Babyrousa babyrussa Babirusa Vv Ss
Choeropsis liberiensis Pygmy Hippopotamus Vv Ss
Vicugna vicugna Vicuna Vv Ss
Dama dama mesopotamica Mesopotamian Fallow Deer E Ss
Cervus duvauceli Barasingha E Ss
Cervus eldi Eld’s Deer Vv $s
Elaphurus davidianus Pere David's Deer E R
Ozotoceros bezoarticus Pampas Deer - Ss
Pudu pudu Pudu - Ss
Okapia johnstoni Okapi - Ss
Tragelaphus euryceros Bongo - Ss
Taurotragus derbianus gigas Giant Eland - Ss
Bubalus (Anoa) depressicornis Lowland Anoa E Ss
Bos gaurus Gaur Vv Ss
Bos javanicus Banteng Vv Ss
Bison bison athabascae Wood Bison - R
Bison bonasus European Bison Vv R
Kobus leche Lechwe Vv Ss
Oryx leucoryx Arabian Oryx E Ss
Addax nasomaculatus Addax E Ss
Gazella cuvieri Cuvier’s Gazelle E ES)
Gazella dama mhorr Mhorr Gazelle E! Ss
Gazella dorcas neglecta Dorcas Gazelle v' Ss
Gazella leptoceros Slender-horned Gazelle E s
Capricornis crispus Japanese Serow - Ss
Ovibos moschatus Musk Ox - Ss
Budorcas taxicolor Takin - Ss
Ammotragus lervia sahariensis Barbary Sheep Vv! Ss
Note: ' category given is that of the whole species; * only some subspecies are included in the studbook; * taxa whose status is under review;
** category refers to A. owenii, the other two species are not threatened.
570
Oryx Project’ was launched by the Sultan of Oman, with
the aim of re-establishing a wild population. In 1980 the
first oryx were returned to Oman for acclimatisation and
eventual reintroduction at Yalooni in the Jiddat-al-Harasis.
In 1982, the first herd of 10 was released from the 1km?
pre-release enclosure into the wild. Further releases were
made in 1984, 1988 and 1989. Numbers increased steadily
and by 1990 there were 109 free-ranging oryx, of which
80% were wild-born, occupying an unrestricted known
range of more than 10,000km?* (Spalton, 1990). Numbers
peaked at 126 in 1991, but a succession of severe drought
years started to produce high mortality. Of the 15 calves
born in 1991, 10 had died by January 1992 and 2 had been
taken into captivity for hand-rearing, leaving the wild
population at 115 (Spalton, pers. comm.). Further releases
are planned to reinforce the wild population
demographically and genetically.
For the first few years of the programme all released
individuals were monitored closely by a force of locally-
recruited Harasis rangers, using radio-tracking equipment
and continuous surveillance from 4-wheel drive vehicles.
Now that numbers have increased, only a selected 40 or so
individuals are monitored. All the oryx are protected from
poaching by strict legislation enforced by the rangers.
Reintroductions from Arizona were hampered by quarantine
restrictions occasioned by the disease blue tongue, which is
endemic in the USA but absent from Oman. Many captive
oryx populations in the Middle East also suffer from
tuberculosis. Proper veterinary procedures were therefore
observed at all stages of the project.
It is estimated that the Yalooni area could eventually
support 200-300 oryx, but competition with increasingly
large herds of domestic livestock is beginning to cause
problems. Agreement has therefore been reached with the
local tribesmen not to graze their herds within a certain
distance of the release site. Nevertheless, with continued
sound managementand effective protection - the keys to the
success of this project so far - the future of the reintroduced
Arabian Oryx at Yalooni seems now to be assured.
The Arabian Oryx reintroduction programme serves to
demonstrate that such projects require the long-term
commitment of substantial amounts of funding and
manpower if they are to succeed. As such, they will of
necessity be confined to a handful of species in the
foreseeable future, and their contribution to the maintenance
of biodiversity will remain very limited.
EX SITU CONSERVATION OF ANIMAL GENETIC
RESOURCES
International efforts to conserve animal species and thereby
preserve animal genetic resources are concerned either with
domesticated or with wild species. At the international level
few programmes attempt to conserve both domestic and
wild species of animals and there is very little interaction
between the two areas.
In the past there has been much less concern over the loss
of genetic diversity in agricultural animals (see Chapter 26)
than for agricultural plants. Consequently, there have only
571
Current Practices in Conservation
been limited attempts to conserve biological diversity in this
area and no programme like the IBPGR presently exists for
animals.
FAO in conjunction with UNEP launched a pilot
programme in 1973 to conserve animal genetic resources.
Initial efforts focused on developing a list of endangered
breeds and of those with economic potential (and to this
extent the remit of this programme was wider than simply
focusing on agricultural animals). In 1980 the FAO and
UNEP called for this programme to be extended and set out
requirements for creating "a supranational infrastructure for
animal breeding and genetics". These requirements covered
a range of efforts to develop animal genetic resources and
amongst other things included guidelines to develop:
databanks for animal genetic resources which would also
identify endangered breeds, gene banks to store semen and
embryos of endangered breeds; training of scientists and
administrators in genetic resource management.
The programme was developed further by a subsequent
Expert Consultation in 1982. This resulted in the FAO
launching its Animal Genetic Resources Programme in
1982. This programme was funded jointly by the FAO and
UNEP and has concentrated on developing methodologies
for a global programme for animal genetic resources. The
work of this programme was published through the FAO
Animal Production and Health Series and included studies
on breed descriptors and databank methodology, the
evolution of cryopreservation and in situ storage of animal
genetic resources.
A five-year programme has recently been proposed by FAO
in which a set of practical field orientated activities will be
carried out. The main features of this programme are: the
preparation of a global inventory of animal genetic
resources; the creation of a ‘World Watch List’ to identify
endangered breeds; breed preservation strategies and
development programmes; development of gene technology
to characterise animal biodiversity and development of a
framework of international undertakings to guide access to
and use of animal genetic resources. This programme
illustrates the growing importance given to this previously
neglected area of biodiversity conservation.
EX SITU CONSERVATION OF MICROBIAL
DIVERSITY
Despite the important role that microbial diversity plays, its
collection and management in the past has been carried out
with a minimum of resources and on an ad hoc basis with
little coordination within a country, let alone on an
international scale. Collections of permanently preserved
living cultures of microorganisms are the microbiologists’
equivalent of botanic gardens, seed banks, zoos, and
aquaria. Such collections are of especial importance to
microbiologists as they are often the only readily available
source of particular organisms required for research and
assessment for exploitation. The reisolation or rediscovery
in nature of desired species is often a matter of chance
alone, and culture collections are thus the essential
mechanism by which the earth’s microbial diversity is made
available to man.
3. Conservation and Management of Biodiversity
One of the first formal attempts to coordinate the
managementof microbial resources on an international level
was the establishment of a directory of institutions
maintaining microbial culture collections. This is now
carried out by the World Federation for Culture Collections
through the World Data Centre under the auspices of
UNESCO, WHO and CSIRO. The latest listing issued by
the WDC details 345 culture collections distributed through
55 countries (Takishima et al., 1989; Table 34.18).
However, many of these collections maintain only a limited
number of strains (mostly under 1,000) and are narrowly
focused (e.g. only plant pathogenic bacteria, or Rhizobium,
or human pathogens).
In 1975 UNEP, the International Cell Research
Organisation and UNESCO jointly called for the
establishment of a worldwide network of culture collections,
and by 1992 there were 16 such collections (Table 34.19).
These collections are known as microbiological resource
centres (MIRCEN). The purpose of these MIRCENs is to
develop and enhance the worldwide network of regional and
inter-regional laboratories. Through this network it is hoped
that a base of knowledge in microbiology will be developed
to support biotechnology in the developing and the
developed world. Activities of MIRCENSs typically include
collection, maintenance, testing and distribution of
microbes, and training of personnel. Though each MIRCEN
works according to its own set of priorities, they share a
common goal of working together to strengthen the network
and advance knowledge in the area. MIRCENs provide the
incentives to develop and maintain microbial collections in
support of national programmes. They also offer a
framework that could provide a secure custodial system for
national and international microbial resources.
Table 34.18 Numbers of collections of living cultures of microorganisms registered
with the World Data Centre on microorganisms
ASIA
China
Hong Kong 1
India
Indonesia
Iran
Israel
Japan
Jordan
Korea
Malaysia
Philippines
Singapore
Sri Lanka
Taiwan
Thailand
Turkey
=
=
i
—---ANAW— -—- AND
USSR (former)
~
EUROPE
Austria
Belgium
Bulgaria
Czechoslovakia 1
Denmark
Finland
France
Germany
Greece
Hungary
Ireland
Italy
Netherlands
Norway
Poland
==
MAN ODADANN WANN — O WW =
EUROPE (continued)
Portugal
Romania
Spain
Sweden
Switzerland
United Kingdom
Yugoslavia
=a 0.) — =
NORTH AND CENTRAL AMERICA
Canada
Guatemala 1
Mexico
USA
SOUTH AMERICA
Argentina
Brazil 1
Chile
Colombia
Venezuela
OCEANIA
Australia
New Zealand
Papua New Guinea 1
AFRICA
Egypt
Kenya
Nigeria
Senegal
South Africa
Uganda
Zimbabwe
LAY pat AC ta Chee er |
Source: Takishima, Y. ef al. 1989. Guide to World Data Centre on Microorganisms with a List of Culture Collections in the World. World Data
Centre on Microorganisms, Saitama.
572
Current Practices in Conservation
Table 34.19 Microbial resource centres (MIRCENs) recognised by UNESCO
Biotechnology MIRCENs
Ain Shams University, Faculty of Agriculture, Shobra-Khaima,
Cairo, Arab Republic of Egypt
Applied Research Division, Central American Research Institute
for Industry (ICAITI), Ave, La Reforma 4-47 Zone 10, Apdo
Postal 1552, Guatemala
Institutet, Fack,
Department of Bacteriology, Karolinska
S-10401 Stockholm, Sweden
Fermentation, Food and Waste Recycling MIRCEN, Thailand
Institute of Scientific and Technological Research, 196
Phahonyothin Road, Bangken, Bangkok 9, Thailand
Fermentation Technology MIRCEN, ICME, University of Osaka,
Suita-shi 656, Osaka, Japan
Institute for Biotechnological Studies, Research and
Development Centre, University of Kent, Canterbury CT2 7TD,
UK
Marine Biotechnology MIRCEN, Department of Microbiology,
University of Maryland, College Park Campus, Maryland
207742, USA
Mycology MIRCEN, International Mycological Institute, Ferry
Lane, Kew, Surrey TW9 3AF, UK
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575
Current Practices in Conservation
Authors as follows: In situ conservation of threatened
plants, Hugh Synge, In situ conservation of crops, G. Brent
Ingram and Alix Flavelle; Ex situ conservation of plants,
E.A. Leadlay and P.S. Wyse Jackson; IARCs, Sam
Johnston; Plant reintroductions, Mike Maunder. Remaining
material by Martin Jenkins and WCMC staff.
3. Conservation and Management of Biodiversity
35.
BACKGROUND
As developing countries have come to realise the economic
value of their biodiversity, so the political considerations
surrounding its utilisation have become more complex. The
direct commodity values of exploited biological resources,
such as tropical hardwoods and fisheries, are well
established, as are the indirect benefits from tourism and
game viewing; what has changed is the recognition of the
value of biodiversity as a genetic resource available for
commercial exploitation through biotechnology.
Developing countries are now demanding a greater share of
the economic benefits arising from the use of resources
within their boundaries which until now have mainly
accrued to the industrial countries with the technological
capability to exploit them.
At the same time, the developed world has become
increasingly apprehensive about the accelerating rate of loss
of biodiversity and its global consequences (of tropical
deforestation upon global climate change, for example).
Developed countries want to see the use of biological
resources placed on a sustainable basis, and are linking
their overseas development assistance to this tenet, which
often clashes with the sovereign rights of developing
countries to manage their resources as they deem best on
behalf of their citizens. In addition, the developed countries
have until recently been able to exploit the genetic resources
of tropical countries for agricultural and pharmacological
advantage at little cost, but have now become concerned
both about the continued erosion of these resources and the
increasing restrictions developing countries are placing on
their use.
These parallel concerns about the exploitation of biological
resources expressed by both the industrialised and under-
industrialised countries have led to the negotiations for a
Convention on Biological Diversity.
Because of the very different interests and expectations of
producer and consumer nations, these negotiations have
become increasingly polarised, with little apparent
willingness to compromise. On the one hand, the developed
countries as consumers of biological resources are
concerned about the sustainability of supplies and continued
unrestricted access to genetic materials, whereas the
developing countries as producer nations are more
concerned with the transfer of the biotechnology to enable
them to develop their resources more effectively for
themselves, and the equitable distribution of benefits arising
from the use of their resources. Redistribution of these
benefits must provide the economic incentive to reinforce
the conservation of biodiversity throughout the developing
world. The obvious difficulties of achieving such
redistribution have greatly retarded progress.
The effects of this increased polarity are exacerbated by
basing the negotiations on the practice of unanimity rather
than consensus, so that a single country with a strongly held
position can insist on alternative wording being included in
the text. The result is a plethora of square brackets
576
THE CONVENTION ON BIOLOGICAL DIVERSITY
including such superficial niceties as "[each Contracting
Party will ....] or [the Contracting Parties shall ....]".
THE BIODIVERSITY CONVENTION
The origin of the convention negotiations goes back to the
initial drafts prepared in 1987 by IUCN in response to a
Resolution adopted at its 16th General Assembly. The
IUCN prototype was a relatively simple document that
focused on measures to reinforce the conservation of
biodiversity in situ through the provision of economic
incentives based on sustainable use. Its main breakthrough,
apart from galvanising activity, was the recognition of the
rights of the producer countries to share equally in the
benefits of their resource use: new, innovative, funding
mechanisms, such as import duties, trade tariffs, and
royalty payments on the sale of commodities incorporating
products of the biological resources of other countries were
proposed.
In 1988, IUCN circulated a comprehensive draft amongst
the participating countries at the UNEP Governing Council.
This stimulated extensive discussion, resulting in acceptance
of the need for an international convention which UNEP, as
the appropriate inter-governmental agency, was instructed
to pursue. The relatively narrow focus envisaged by IUCN
was then expanded to include ex situ conservation, land
races and the wild relatives of commercial crop varieties,
access to technologies and scientific skills by developing
countries, and the transfer of biotechnologies for developing
countries to exploit their own genetic resources. Formal
negotiations commenced in November 1990 with the first
session of the Ad Hoc Working Group of Legal and
Technical Experts, followed by five sessions of the
Intergovernmental Negotiating Committee (INC) for a
Convention on Biological Diversity. Each INC meeting has
involved some 75 countries and lasted for eight days,
representing a substantial investment of time and funding.
As with the Convention on Climate Change, the
Biodiversity Convention has become a key component of
the UN Conference on Environment and Development
(UNCED), with the expectation of a formal instrument
being ready for signature at the time of the conference in
Brazil. However, with many of the most substantive issues
still to be agreed, it is probable that only a framework
convention will be ready for Rio, with the more contentious
articles being negotiated as protocols to the convention
according to a schedule to be decided at UNCED.
The effects of these delays will obviously influence the
timetable for the convention coming into operation. As with
other international legal instruments, countries must first
sign and then subsequently ratify the convention, and it is
only after the twentieth country has acceded (and this
number is still subject to debate) that it comes into force.
With ratification likely to be delayed until the protocols
have been negotiated, it may be five years or more until the
convention becomes operational. The non-governmental
organisations are already voicing their concerns about the
continued loss of biodiversity before the convention comes
into force.
Table 35.1
fe Objectives
2. Use of Terms for the Purpose of this Convention
oe Fundamental Principles
4. General Obligations
Se Implementation Measures
[ 5 bis Identification and Monitoring]
6. In situ Conservation
Ts Ex situ Conservation
8. Sustainable Use of Components of Biological
Diversity
[ 8 bis Incentive Measures]
ch Research and Training
10. Public Education and Awareness
Ais Impact Assessment
Duti2. Surveys and Inventories]
Dass Global Lists]
[ 14. Access to Genetic Material]
[ 14 bis Traditional Indigenous and Local Knowledge]
Wen Access to Technology
[ 15 bis Exchange of Information]
16. Transfer of Technology
17. Technical and Scientific Cooperation
17 bis Handling of Biotechnology and Distribution of its
Benefits
The Convention on Biological Diversity
Articles of the draft Convention on Biological Diversity
18. Financial Needs and Means
Ue Financial Mechanisms
20. Relationship with other International Conventions
2s Conference of the Parties
[ 22. Procedures for Global Lists]
23. Secretariat
24. Science and Technology Committee
25. Reports
26. Operational Cost
ie Settlement of Disputes
28. Adoption of Protocols
29. Amendment of the Convention or Protocols
30. Adoption and Amendment of Annexes
31. Right to Vote
32. Relationship between the Convention and its
Protocols
33. Signature
34. Ratification, Acceptance or Approval
Soe Accession
36. Entry into Force
37. Reservations
38. Withdrawals
39. Depository
40. Authentic Text
Notes: This table reflects the fourth revised draft of the Convention (February 1992).
[ ] = Articles in square brackets are still subject to
negotiation and may be dropped in their entirety. Most of the operational articles include disputed clauses, or text within clauses, in square brackets.
The contents of the convention
The contents of the draft convention are outlined in Table
35.1. Although the exact measures to be incorporated in the
articles are still to be decided - and at the opening of the
fourth session of the INC in February 1992 (only four
months before UNCED) there were still over 350 disputed
sections of text - certain features have already begun to
emerge (at the time of writing this review the text of the
convention was still under negotiation; this analysis is based
on the provisions of the Fourth Revised Draft, including the
compromise formulations proposed by the Executive
Director of UNEP, which was presented to the Fourth
Session of the INC, 6-15 February 1992).
The Objective (Article 1) of the convention is "to conserve
the maximum possible biological diversity for the benefit of
present and future generations and for its intrinsic value",
which is to be achieved by:
© ensuring that the use of biological resources is
sustainable
providing adequate, new and additional funding for
developing countries to facilitate the conservation and
rational use of their resources
taking account of the need to share costs and benefits
between developed and developing countries
providing economic and legal conditions favourable for
the transfer of technology necessary to accomplish the
objectives of this convention
providing fair sharing of the benefits of research in
biotechnology arising from the conservation of biological
diversity.
The Fundamental Principles (Article 3) are extremely
broad: they affirm that the conservation of biodiversity is a
common concern of all people, but also stress the
577
responsibility of states in exercising their sovereign rights
to ensure that their biological resources are developed in a
sustainable way. Emphasis is also given to the in situ
conservation of ecosystems and natural habitats, whilst ex
situ measures should preferably be undertaken in the
country of origin. An important recognition is that lack of
scientific certainty should not be used as a reason for
postponing actions to avoid or minimise threats to
biodiversity. Regarding sources of finance, countries
benefiting most from the exploitation of biodiversity should
contribute most to its conservation. The practices and
experience of indigenous peoples in using biological
resources should be recognised and rewarded.
The General Obligations (Article 4) still have much disputed
text, but call on each Contracting Party to take all measures
at its disposal, including national plans, policies and
legislation, both individually and cooperatively, to conserve
the maximum possible biological diversity within its
national jurisdiction.
The Implementation Measures (Article 5) envisage the
development of national strategies and programmes for the
conservation and sustainable use of biological diversity,
including the establishment of national bodies to implement
the provisions of the Convention.
The next ten articles (Articles 5 bis to 13 inclusive)
comprise the conservation core of the convention. They
cover such issues as in situ and ex situ conservation,
inventory and monitoring, research and training, public
education, and sustainable use. The obligations on countries
have been weakened by obstruction from the USA and some
of the G77 states (the group of 77 non-aligned developing
countries), but still include a number of positive features.
For example, countries are called upon to: establish
protected areas in locations requiring special conservation
3. Conservation and Management of Biodiversity
measures, including wildlife corridors; restore degraded
ecosystems and habitats; eradicate alien species that threaten
natural habitats; and to introduce legislation for the
protection of threatened species, populations and varieties.
Parties are also expected to undertake national surveys of
their biodiversity and to maintain databases of their
resources, linked into a global network. However, the
convention makes it quite clear that the implementation of
these obligations by developing countries is subject to the
provision of new and additional financial and technical
resources.
There then follow a series of articles (numbers 14 - 17 bis)
that deal with access to genetic resources and the transfer of
the technologies and scientific skills appropriate for their
exploitation. The thrust of this section is that countries
should refrain from imposing restrictions on the availability
of wild genetic materials, such as breeders’ or farmers’
rights, but that preferential access to the research results or
benefits arising from the use of genetic materials should be
granted to the country of origin. At the same time, parties
should undertake to provide, on mutually agreed terms,
technologies appropriate to the conservation and sustainable
use of biodiversity.
Aside from Articles 18 and 19 (see below), the remaining
articles relate to the procedures for the establishment and
operation of the convention and its protocols, and are
therefore less contentious. This administrative machinery is
essential for the development of an effective convention,
and the procedures that have now been agreed are the most
sophisticated yet seen in an environmental treaty.
Some contentious issues
Articles 18 and 19 cover the key issues of financial needs
and mechanisms, upon which the viability of the whole
convention depends. There is a general expectation that the
developed countries must provide "adequate new and
additional financial resources to enable developing countries
to meet the agreed incremental costs to them of fulfilling
their obligations under the Convention". There is also
widespread acceptance for the establishment of a Biological
Diversity Fund for developing countries to implement their
obligations, but a difference of opinion about how this fund
should be administered. One option is to create a ‘window
account’ in the Global Environmental Facility (GEF)
specifically for the convention, although the criteria for
allocating these funds would be determined by the
Conference of the Parties through a Science and Technology
Committee, rather than by the three agencies in the GEF
(see Chapter 32 for an outline of the GEF).
Whatever administrative mechanism is adopted, the purpose
of this funding will be to empower developing countries to
meet the scientific, economic and institutional requirements
of the convention. It is not envisaged that the Biological
Diversity Fund should provide the conduit for the economic
incentives that developing countries may need to reinforce
their conservation programmes: these must be derived from
standard commercial practices arising from the use of
biological resources negotiated by national governments.
In addition to the central issue of the financial provisions,
the other contentious issues where substantive differences of
opinion still need to be resolved include:
e the granting of access to genetic resources and the
conditions pertaining to their use
© equitable distribution of benefits arising from the use of
genetic resources between the exploiting country and the
country of origin
© provisions for biotechnology safety relating to the
introduction of genetically modified organisms
e fair and favourable conditions for access to and transfer
of technology
® commercial patents and intellectual property rights
relating to the transfer of biotechnological processes and
genetic manipulation procedures
e the global lists of species and sites
Article 13 calls for a Global List of Biogeographic Areas of
Particular Importance for the Conservation of Biological
Diversity and a second Global List of Species Threatened
with Extinction on a Global Level, but this proposal has run
into opposition. Some developing countries are against lists
because of the burden they would impose if species
recovery plans and site management plans had to be
implemented; also, global lists could be seen to conflict
with the rights of national sovereignty if designations were
imposed on countries without their agreement.
On the positive side, lists would focus world attention on
the sites and species of global conservation concern, and
would help identify priorities for funding. The lists would
represent a tangible output from the convention and provide
a vehicle for a concerted conservation effort involving the
non-governmental organisations. It is no coincidence that
the more effective conservation conventions, such as The
World Heritage Convention (WHC), Ramsar and CITES,
all have lists at their core. The non-governmental agencies
are lobbying hard for the retention of lists on the grounds
that their removal would greatly dilute the conservation
provisions (see International obligations, Chapter 31).
The key role of an active administrative structure
With many of the substantive provisions of the Convention
on Biological Diversity still undecided, the sophisticated
administrative structure already agreed will be vital in
developing the convention to a stage where it is a truly
effective international instrument; this structure is in fact a
major achievement of the negotiations to date.
The vital role that an active administration plays in
developing a framework convention into an effective
international instrument is illustrated by existing
conventions. The success of both CITES and the WHC is
largely due to these conventions having active and well-
financed administrative structures. The comparative failure
of the Bonn Convention or the Western Hemisphere
Convention is in part attributable to the absence of such
structures. The essential features include: a well-financed
secretariat; an independent scientific committee;
requirements for regular meetings of the parties and regular
reporting by them to the secretariat; the involvement of
outside parties (such as NGOs) in the regular meetings of
the parties; and the obligation to establish or designate a
national or local authority to deal with implementing the
obligations of the convention.
The effectiveness of these measures arises from the fact that
they keep the key issues in the public arena and on the
political agenda, thereby working against political and
administrative inertia. They also provide a catalyst for
development of the broad objectives which a framework
convention largely comprises into specific obligations which
have some impact on the conduct of the Parties.
The Convention on Biological Diversity has many of these
features. The convention establishes a Secretariat (Article
23) to arrange and coordinate meetings of the Parties; to
assist the scientific committee in its work; and to maintain
the global lists, if these are to be included. The functions of
this body are to be carried out by an existing international
organisation, to be decided at the first conference of the
Parties, but which in the interim will be the responsibility
of UNEP. The convention establishes a scientific committee
which is called the Scientific and Technology Committee;
its role will be to provide scientific and technological advice
as required for the implementation of the convention. There
are extensive reporting requirements and meetings of the
Parties are to be held at regular intervals to be decided at
the first conference of the Parties. Non-governmental
organisations are eligible to attend these conferences
provided they have informed the secretariat and not more
than one third of the Parties object to their presence.
THE BIODIVERSITY COUNTRY STUDIES AND
UNMET FINANCIAL NEEDS
The conservation element of the convention focuses initially
on the gathering of information through national surveys
and inventories, then moves on to address the benefits
arising from the sustainable use of biodiversity. This
information collecting exercise is to be undertaken by
Country Studies detailing what is currently known about the
status, threats, costs and benefits of biodiversity in each
country (Table 35.2). The Country Studies will then form
the basis for the development of the national plans for the
conservation and sustainable use of biodiversity called for
under the Implementation Measures.
At the same time, the INC needs to quantify what order of
magnitude of new and additional financial resources will be
required for the Biological Diversity Fund to finance the
implementation of the measures in the Convention by
developing countries. The Country Studies were therefore
charged with calculating the unmet financial needs of each
country undertaking a survey from which the total financial
requirements of the Convention could be estimated.
With the coordination of UNEP, GEF funding, and the
World Conservation Monitoring Centre playing a catalytic
role, some 14 countries, of which 11 have reported, are
undertaking Country Studies. A methodology for
completing the studies was prepared by UNEP and has four
main components:
review of the status of the biological resources
identification of the measures necessary for effective
conservation and sustainable use of these resources
determination of the costs and benefits of implementing
these measures
estimation of the current unmet financial needs
579
The Convention on Biological Diversity
This process was expected to furnish many new data on the
status and economics of biodiversity conservation and
utilisation (Table 35.2). In practice, the methodology has
proved to be over-ambitious so that even developed
countries have had problems implementing it, although
extensive new data on biological resources have been
forthcoming. Of equal importance has been the recognition
of the gaps in the information-base, particularly
microorganisms, invertebrates and lower plants.
Table 35.3 shows the estimated unmet financial costs of the
ten reporting countries. The substantial variation in the
annual needs, ranging from US$1,590/km? for Costa Rica
to US$64/km*? for Kenya, reflects more the lack of
standardisation in the estimation than real differences in
financial requirements. Clearly these needs will vary
significantly between countries - for example, countries
with a sound infrastructure for biodiversity conservation
will require fewer funds to implement measures in the
convention than those with a neglected infrastructure.
Extrapolating from these figures of unmet costs, a number
of estimates have been made, using different methods, to
quantify the total financial resources required by all
developing countries to implement the convention (Table
35.4) (UNEP, 1992). The average of these estimates is
around US$20 billion/annum. Although no more than
indicative of the order of magnitude, this estimate does
suggest that substantial amounts of additional funds will
have to be transferred to developing countries if they are to
meet their obligations under the convention.
The current level of overseas development assistance
available to developing countries for the conservation of
biological diversity is estimated at US$228 million, of
which US$170 million is derived from bilateral aid and
US$58 million from multilateral sources (UNEP, 1991). A
ten-fold increase in commitment from the donor countries
is therefore required. Considering the current apprehensions
being expressed by the developed countries about the likely
levels of extra funding that the convention will need, it is
politically inconceivable in the short-term that additional
funding of this magnitude will be forthcoming. Although
US$20 billion/annum in absolute terms is a substantial sum,
representing about 27% of the total overseas aid budget, it
is put into a realistic context by comparison with the $245
billion spend each year by the OECD countries on their
own agricultural support programmes, which are themselves
ultimately dependentupon biodiversity. With global military
budgets at some US$980 billion in 1990, or US$185 for
every person on the planet, a peace dividend from the
cessation of the cold war of only 2% would seem a modest
amount to save the diversity of life on earth.
Based on a pragmatic assessment of what developed
countries are likely to find acceptable, UNEP is proposing
the establishment of a roll-over mechanism based on the
GEF to provide interim funding for the convention. This
would accelerate once the institutional and human capacity
were in place. UNEP is proposing an increase of US$500
million/annum over the next five years, raising the current
flow of funding through the GEF from around US$100
million/annum to US$600 million/annum by 1997, and
thereafter accelerating to US$850 million/annum by the end
3. Conservation and Management of Biodiversity
Table 35.2 Information to be generated by Biodiversity Country Studies using the
UNEP methodology of reporting
Annex | Global and National Biodiversity Status
A. Species diversity data
B. Species ecological status over time
C. Habitat/ecosystems diversity
D. Habitat/ecosystem status and percent change over the past 10-20 years
E. Areas of high species endemism
F. Significant changes in populations of species of national importance over the past 10 years
G. National parks/nature reserves/gazetted forests and other protected sites
H. Additional national biotic communities/biogeographic provinces currently not protected
|. Private wildlife sanctuaries
J. Status of national ex situ conservation facilities
K. Species in national ex situ conservation facilities
Annex II Essential Planetary Services Provided by Major Taxonomic Groups of Organisms
Annex Ill Categories of Value Assigned to Biological Diversity
Annex |V_— Sites and Species of Significance for Conservation
Annex V Measures to be Implemented to Achieve Desired Level of Conservation
Annex VI Measures to be Undertaken for Effective Conservation and Rational Use of Biological Diversity
Annex Vil Calculating Costs and Benefits Associated with the Implementation of Identified Measures for Conservation and
Sustainable Use of Biological Diversity
Annex IX Current Multilateral, Bilateral and National Financial support for Biodiversity Conservation and Unmet Funding Needs
in Respect of Identified Priority Areas
Annex X Summary of Costs, Benefits and Unmet Needs of Biodiversity Conservation
Table 35.3. Unmet financial needs of countries to conserve their biodiversity
COUNTRY TOTAL ANNUAL COSTS UNMET ANNUAL COSTS
TOTAL PER KM?*
US$ million/year US$ million/year US$ million/year
ASIA
Indonesia 290 231 120
Malaysia x x »4
Thailand 120 60 116
EUROPE
Germany 1,200 950 2,662
Poland 800 100 320
SOUTH AMERICA
Guyana -- -- -
Peru -- -- --
NORTH AND CENTRAL AMERICA
Bahamas 110 84 **6,058
Costa Rica 100 81 1,590
Canada 2,686 986 99
OCEANIA
Australia -- -- --
AFRICA
Kenya 160 37 64
Nigeria 593 325 352
Uganda 70 58 245
Source: Data derived from Country Study reports.
Notes: X relevant economic data not supplied; -- no Country Study Report submitted; * terrestrial land area only; ** excludes area of marine
habitats.
580
The Convention on Biological Diversity
Table 35.4 Estimates of the total unmet financial needs per annum of all
developing countries to implement the measures in the Convention
on Biological Diversity
METHOD OF ESTIMATION
TOTAL UNMET FINANCIAL NEEDS
USS billion/annum
A. Extrapolation on the basis of unmet needs of developing countries 8.45
adjusted for biodiversity richness and country size
B. Extrapolation on the basis of percentage GDP (0.5%) as desirable 21.13 to 42.25
expenditure for biodiversity conservation and sustainable use
Cc. Extrapolation on the basis of the number of sites and national 42.0
protected areas as a percentage of the global total
D. Extrapolation on the basis of species diversity in each country as a 0.68 to 15.8
percentage of global total
E. Extrapolation on the basis of categorising countries by their Wile
biodiversity richness and their in-situ conservation infrastructure (the
WCMC method)
of the century. Although welcome, this level of funding is
still far short of the minimum requirement estimated from
the Country Studies.
FUTURE DATA NEEDS: NETWORKING AND
GLOBAL MONITORING
The preparation of the Country Studies has proved to be a
valuable mechanism for setting a country on course towards
a better understanding of its biodiversity and a more
rational use of its resources. The process has necessitated
the establishment in each country of National Biodiversity
Units (NBUs) to serve as coordinating centres for the
gathering of data on the status, utilisation, and economic
values of biodiversity. The concept of accounting the costs
and benefits of biodiversity conservation and use has been
introduced, and appreciation of the costs of inaction in
terms of lost benefits if no action is taken to conserve
biodiversity, has been accepted.
The Country Studies exercise and the resultant NBUs will
provide a useful foundation upon which to build the human
and institutional capacities for improved conservation
practice. Already further studies are being planned for some
of the most biologically rich countries such as Brazil,
Colombia, Mexico, Madagascar, Zaire and Papua New
Guinea, as well as countries such as Angola and
Mozambique where the conservation infrastructure requires
rebuilding. However, expanding the programme too rapidly
will divert the limited GEF funds away from the priority
activities of consolidating the results of the first tranche of
reporting countries and of providing long-term support for
those NBUs that have already proved their worth. If the
programme is to advance it must first revise the
methodology to produce a more robust system for
estimating economic costs and benefits which can be applied
realistically in developing countries. It must be accepted
that quantifying the existence value of a threatened species
or the service value of a wooded watershed to secure a
constant water supply involves an element of subjective
value judgement, but guidelines are needed so that estimates
can be standardised between countries.
581
In addition, the NBUs should be further developed and
strengthened into National Biodiversity Monitoring Centres
responsible for the gathering and analysis of data at the
country level. Such monitoring centres should then be
linked into a global biodiversity information network which
can be progressively expanded with each subsequent round
of Country Studies. This proposal closely mirrors the key
recommendation of the Global Biodiversity Strategy for the
establishment of an Early Warning Network to monitor
potential threats to biodiversity (Table 35.5)
(WRI/IUCN/UNEP, 1992). The purpose of this Network is
to provide a swift response to the emergence of new threats
through the rapid mobilisation of information.
The best sources of early warning information are
scientists, non-governmental organisations, and enforcement
authorities working in the field. If they can be linked into
an in-country network of data sources feeding their
information into a National Biodiversity Monitoring Centre,
which in turn is linked into a global network, then a
mechanism can be developed to mobilise this information
rapidly. The parameters that a national centre should
monitor for early warning purposes must include not only
direct threats but also political, legal and economic changes
that could have indirect effects on biodiversity. A set of
such parameters is presented in Table 35.6.
Incorporating biodiversity conservation into national policies
and planning (Action 5 of Table 35.5) can help countries
define and articulate their environment and development
goals. A minimum set of biodiversity indicators that must
be included within the monitoring programmes of a national
data centre, and which provide the basic information needs
for national and international policy-makers is presented in
Table 35.7. This dataset provides a matrix combining the
major conservation concerns with a working set of
indicators that can be used to assess long-term trends in the
conservation of biodiversity.
The limiting factor in such programmes for assessing
biodiversity conservation trends and goals is the availability
3. Conservation and Management of Biodiversity
Table 35.5 International actions to conserve the world’s biological diversity as
recommended by the Global Biodiversity Strategy
Action 1. Adopt in 1992 the International Convention on Biological Diversity
2. Adopt in the General Assembly of the United Nations, a resolution designating 1994-2003 the International Biodiversity
Decade
3. Establish a mechanism, such as an International Panel on Biodiversity Conservation, preferably within the Convention
on Biological Diversity, including scientists, non-governmental organisations and policy-makers to provide guidance
on priorities for the protection, understanding, and sustainable and equitable use of biodiversity
4. Establish an Early Warning Network, linked to the Convention on Biological Diversity, to monitor potential threats to
biodiversity and mobilize remedial action
2h Integrate biodiversity conservation into national planning processes
Source: WRI/IUCN/UNEP 1992. Global Biodiversity Strategy.
Table 35.6 Parameters that an Early Warning Network must monitor at the country
level
1. Traditional crop or livestock varieties threatened by planned development projects or the introduction of new varieties
2. Increasing genetic uniformity of crops
3. Natural ecosystems subjected to new inappropriate management practices, human encroachment, or unsustainable exploitation
4. Protected areas in urgent need of financial, technical, or other support
5. Accelerating habitat loss
6. Evidence of the over-exploitation of species
7. Introductions of exotic species
8. Genebank facilities with germplasm at risk due to lack of funding for recurring costs
9. Climatic threats to biodiversity - including desertification, floods, drought, and global warming
14.
15.
Source: WRI/IUCN/UNEP 1992. Global Biodiversity Strategy.
and reliability of the data. In collaboration with WCMC,
Reid et al. (in prep.) reviewed the availability, coverage
and quality of the data needed for assessment of the
indicators presented in Table 35.7. The conclusions make
depressing reading: although the coverage at the country
level for mammals and birds is reasonable, for most other
species the data are lacking or, where available, of poor
quality. Time series data are non-existent except for a few
“‘megafauna’ species and for the land-use estimates by FAO.
Of particular concern are the lack of data on genetic
varieties of agricultural crops grown in developing
countries, and the absence of base-line datasets for
monitoring ecosystem changes.
582
. Communities denied access to resources when protected areas are established
. Pollutant discharges presenting immediate threats or chronic pollution that might pose longer-term threats
. Changes to the legislation relating to land and other resource ownership that may disenfranchise local communities
. Changes to national budgets that may affect the allocation of funds for conservation
Political or institutional developments that may influence the infrastructure for effecting conservation
Implementation of obligations undertaken through international conventions
These conclusions emphasise the urgent need to build
monitoring capabilities at the country level. With the
increasing precision of remote sensing techniques and the
advances in information technology, particularly the
application of Geographic Information Systems, the ability
to develop sophisticated monitoring systems is within the
reach of all countries. In an analysis of the relevance of
technology transfer to the conservation of biological
diversity, a recent report (Touche Ross, 1991) showed that
the most appropriate technologies for the management and
utilisation of biodiversity were ‘soft’: that is, information
management, human skills and scientific knowledge rather
than ‘hard’ involving physical plant and equipment.
The Convention on Biological Diversity
Table 35.7. A minimum set of indicators for monitoring biodiversity at the country
level
INDICATOR BIODIVERSITY CONSERVATION CONCERNS
GENETIC SPECIES COMMUNITY
DIVERSITY DIVERSITY DIVERSITY
Wild Species and Genetic Diversity
ile Species richness (number, number per unit area, number per e e®
habitat type)
Pe Species threatened with extinction (number or percent) e e
3. Species threatened with extirpation (number or percent) e e
4. Endemic species (number or percent) e e
= Endemic species threatened with extinction (number or e e
percent)
6. Species risk index e ®
7 Species with stable or increasing populations (number or e e
percent)
8. Species with decreasing populations (number or percent) ® e
oF Threatened species in protected areas (percent) @ ®
10. Endemic species in protected areas (percent) e e
ils Threatened species in ex s/tu collections (percent) ® e
12. Threatened species with viable ex situ populations (percent) e e
13. Species used by local residents (percent) e e
Community Diversity
13. Percent dominated by non-domesticated species e ®
14. Rate of change from dominance of non-domesticated species e e
to domesticated species
15. Percent of area dominated by non-domesticated species e e
occurring in patches greater than 1,000km?
16. Percent of area in strictly protected status e e
Domesticated Species
17. Accessions of crops and livestock in ex situ storage (number) e
18. Accessions regenerated in the past decade (percent) e
19° Number of crops (livestock) grown as percent of number 30 e
years before
20. Number of varieties as percent of number 30 years before e
2. Coefficient of kinship or parentage of crop e
Source: Reid, W.V., McNeely, J.A., Tunstall, D.B. and Bryant, D. (in prep.). World Resources Institute, Washington DC.
The main issue now is not how to monitor but what to
monitor. A minimum set of parameters must be agreed,
along the lines presented in Table 35.6 and 35.7, that
provides a framework for determining conservation
priorities and goals at the country level, that generates the
data necessary to build biodiversity conservation into the
national planning process, and that supplies the early
warning information necessary for the rapid response to
new threats. Such a system will require the standardisation
of species names, habitat classifications and threat
categories so that national data centres can be networked for
the reciprocal exchange of information. As more centres are
established, so the network will grow, enabling regional
assessments to be made of needs, priorities and financial
investments.
This need to build the information capacity as the basis for
decision-making is recognised as the first prescribed action
in the biodiversity proposals for Agenda 21 of UNCED. An
encouraging start has been made with the Country Study
Programme, which must now be expanded through the
Convention on Biological Diversity to develop the human
skills and monitoring capabilities of developing countries.
The long-term goal must be to create a global biodiversity
583
information network, linking the national centres and
mobilising the substantial amounts of data worldwide to
promote a more enlightened conservation and development
practice.
References
Reid, W.V., McNeely, J.A., Tunstall, D.B. and Bryant, D. (in prep.).
World Resources Institute, Washington DC.
Touche Ross 1991. Conservation of Biological Diversity: the role of
technology transfer. A Report for the United Nations Conference
on Environment and Development and the UNEP
Intergovernmental Negotiating Committee for a Convention on
Biological Diversity. Touche Ross and Co., London. 67pp.
UNEP 1991. Guidelines for the Preparation of Country Studies on
Costs, Benefits and Unmet Needs of Biological Diversity
Conservation within the Framework of the Convention on Biological
Diversity. UNEP/Bio.Div./Guidelines. May 1991. 73pp.
UNEP 1992. Biodiversity Country Studies: executive summary. A
Report to the Fifth Session of the Intergovernmental Negotiating
Committee for a Convention on Biological Diversity.
UNEP/Bio. Div./N7-INC.5/May 1992. 6pp. —
WRI/IUCN/UNEP 1992. Global Biodiversity Strategy: guidelines for
action to save, study and use earth’s biotic wealth sustainably and
equitably. World Resources Institute, Washington DC. 243pp.
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GLOSSARY
This highly selective Glossary provides definitions of some
of the less familiar or more technical terms to be found
above.
Anadromous: (of fishes) those which ascend rivers from
the sea in order to spawn.
Anaerobic respiration: liberation of energy by breakdown
of substances not involving consumption of oxygen.
Archaean: belonging to or containing the group of rocks of
the Archaeozoic era (the earlier part of the Precambrian
era), which ended about 2,500 million years ago.
Back-arc basins: ocean floor spreading centres associated
with subduction processes in deep ocean trenches.
Catadromous: (of fishes) those which descend rivers to the
lower or estuarine reaches, or the sea, in order to
spawn.
Chloroplasts: plastids containing chlorophyll, sometimes
with other pigments, found in the cytoplasm of higher
plant cells.
Chromophyte*: a member of the Chromophyta, a major
division of the plant kingdom including most of the algae
characterised by the presence of flagellae on the sexual
spores and the presence of chlorophyll a (but not 5).
Coccoliths: minute mainly marine protoctistan organisms
(formerly treated as protozoa or algae) dating from
Cambrian to modern times, with calcium carbonate ring
or platelet structures, which form substantial chalk
deposits when fossilised.
Conidial: used of fungi producing conidia, that is asexual
spores formed by mitotic divisions; sexual stages are
unknown in many such fungi, which are sometimes
referred to as ‘deuteromycetes’ or ‘imperfect fungi’.
Cyanobacteria: photosynthetic and nitrogen-fixing blue-
green bacteria, formerly generally treated as blue-green
algae but lacking nuclei and therefore regarded as
belonging to the bacterial Kingdom.
Diploid: having two sets of chromosomes in the nucleus of
each somatic cell. Characteristic of most normal
eukaryotic higher organisms.
Ectomycorrhiza: mycorrhiza (q.v.) where the fungal
mycelium is only associated with the first layer of
epidermal cells on the plant root.
Endomycorrhiza: mycorrhiza (q.v.) where the fungal
mycelium penetrates into the cortex of the plant root.
Epedaphon: inhabitants of the soil surface, e.g. most
ground-beetles and scorpions.
Euedaphon: inhabitants of the mineral soil, e.g. most
earthworms, all Symphyla, many mites.
Eukaryote: a cell or organism with a membrane-bounded
nucleus, organelles and chromosomes with histone-
coated DNA.
Germplasm: genetic material, especially its specific
molecular and chemical constitution, that comprises the
physical basis of the inherited qualities of an organism.
Haploid: having the number of chromosomes characteristic
of the gametes for the organism (one set in most
eukaryotic normal higher organisms).
Hemiedaphon: inhabitants of the litter and fermentation
layer, e.g. many woodlice and millipedes.
Heterotrophic heterokonts: filamentous or unicellular
organisms, the sexual spores (zoospores) of which have
two hair-like appendages each of a different structure,
and which also lack chlorophyll and obtain the
carbohydrates they require by parasitising plants or
utilising dead organic materials.
Hexapods: six-footed animals, specifically the insects
(although certain primitive forms are occasionally
excluded from the taxon Insecta and these groups
together are included in the Hexapoda).
Mitochondria: double-membraned organelles in the
cytoplasm of all eukaryotes where the respiratory cycles
occur.
Mollicute: a bacterium-like organism such as Spiroplasma,
lacking an independent wall and always occurring inside
the cells of cellular organisms, particularly insects.
Morphospecies: a group of individuals which are
considered to belong to the same species on
morphological grounds alone.
Mycorrhiza: mutualistic symbiotic associations between
fungi and the roots of green plants, occurring in about
80% of all vascular plants and also certain bryophytes;
the fungi either form nets over the root surfaces or are
mainly confined to a special layer within the root tissues
themselves.
Ocean-floor spreading centres: these occur along the
central axis of most oceans, and are the seismically
active regions where new oceanic crust forms; as new
crust is extruded from below, the older crust is pushed
away from the axis of the ridge.
Organelles: the various inclusions in a cell which have
special functions, e.g. mitochondria and chloroplasts.
Picoplankton: minute algal-like organisms with cells about
2 microns in diameter abundant in the upper layers of
the world’s oceans.
Plastid: a cytoplasmic, pigmented photosynthetic organelle
or its non-photosynthetic derivative.
Primary productivity: the rate of transformation of
chemical or solar energy to biomass. Most primary
production is carried out by plants through
photosynthesis but some bacteria can convert chemical
energy to biomass through chemosynthesis.
Prokaryote: a cell or organism composed of cells lacking
a membrane-bound nucleus, membrane-bound organelles
and histone-coated DNA.
Protoctists*: eukaryotic organisms which are not plants,
animals or fungi, i.e. protozoans and other unicellular
organisms, algae, slime moulds, etc.
rRNA: ribosomal ribonucleic acid - the type of RNA
which, together with proteins, makes up the ribosomes.
Subduction and fracture zones: where spreading ocean
crust impinges against an unyielding continental margin,
an ‘active margin’ forms, with the oceanic crust buckling
downwards (subducting) and being destroyed within the
hot interior of the earth; ocean trenches are formed
along these margins.
Note: * Dependent on classification system used.
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Global Biodiversity: ; re
Status of the Earth’s living resources ds :
Compiled by eek)
. . . WORLD CONSERVATION
World Conservation Monitoring Centre MONITORING CENTRE
Global Biodiversity: Status of the Earth's living resources is the most
comprehensive compendium of conservation information ever publish-
ed. It provides the first systematic report on the status, distribution,
management, and utilisation of the planet’s biological wealth. With
extensive use of tables, graphics and maps, it presents standardised
and comparable data for 205 countries of the world. The report is
divided into three sections:
@ Part 1 documents diversity at the genetic, species and ecosystem
levels, including micro-organisms, species extinctions and rates of
habitat loss.
@ Part 2 covers the benefits and values of biodiversity, focusing on
wildlife utilisation, domestication and the economic valuation of
diversity.
@ Part 3 reviews the conservation and management of biodiversity at
the national and international levels, including i” situ and ex
situ Management, international conventions, and institutional and
financial support.
This authoritative reference-work will be an indispensable source of
factual information for everybody involved in conservation and develop-
ment — politicians, planners, resource managers, aid workers, scien-
tists, journalists, and the concerned general public.
The Global Biodiversity Strategy is a comprehensive set of guidelines
and actions to save, study and utilise the Earth’s biotic wealth. Publish-
ed by the World Resources Institute, IUCN — The World Conserva-
tion Union, and the United Nations Environment Programme, it
prescribes a decade of action to slow the loss of biodiversity by pro-
moting its sustainable and equitable use. G/oba/ Biodiversity: Status
of the Earth’s living resources provides the platform of scientific data
upon which to implement the Strategy and to monitor its effects.
||
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Saving Our Planet
Challenges and hopes
M.K. Tolba
Paperback (0 412 47370 4), 304 pages
The World Environment 1972-1992
Two decades of challenge
Edited by O.A. El-Kholy, M.W. Holdgate, D.F. McMichael,
R.E. Munn and M.K. Tolba
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